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HARVARD  UNIVERSITY 


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LIBRARY 


OF  THE 


Museum  of  Comparative  Zoology 


APR  12  H 


r^r    ' 


BULLETIN  OF  THE  UNIVERSITY  OF  KANSAS 

Vol.  XXIII  DECEMBER,  1922  No.  18 

Science  Bulletin 

Vol.  XIV,  Nos.  1,  2,  3,  4,  5,  6,  7,  8,  9,  10,  11,  12,  13,  14, 
15,  16, 17,  18, 19,  20  and  21 

(Continuation  of  Kansas  University  Quarterly.) 


ENTOMOLOGY  NUMBER  V 


LAWRENCE,  KANSAS 

Published  Semimonthly  from  Januaiy  to  June  and  Monthly  from  July  to 
December,  inclusive,  by  the  University  of  Kansas. 


Entered  as  second-class  matter  December  29,  1910,  at  the  post  oflBce  at  Lawrence,  Kansas, 

under  the  act  of  July  16,  1894. 


9-4522 


NOTICE  TO  EXCHANGES. 

The  attention  of  learned  societies  and  other  institutions  which 
exchange .  s(^ie!ntifi|C  publications  with  the  University  of  Kansas  is 
called  to  the  list  of  publications  of  this  University  on  the  third  and 
fourth  pages  of  the  cover  of  this  issue. 

Those  marked  ''Supply  exhausted"  cannot  be  furnished  at  all ;  as 
far  as  the  supply  permits  the  remaining  numbers  will  gladly  be  fur- 
nished to  any  of  our  exchanges  who  may  need  them  to  complete  their 
files. 

Back  numbers  of  the  Kansas  University  Quarterly,  as  far  as  pos- 
sible, will  be  sent  to  those  of  our  newer  correspondents  who  are  able 
and  willing  to  reciprocate. 


ANNOUNCEMENT. 

The  Kansas  University  Science  Bulletin  (continuation  of  the 
Kansas  University  Quarterly)  is  issued  in  parts  at  irregular  inter- 
vals. Each  volume  contains  from  300  to  400  pages  of  reading-mat- 
ter, with  necessary  illustrations.  Exchanges  with  other  institutions 
and  learned  societies  everywhere  are  solicited.  All  exchanges  should 
be  addressed  to  the  Library  of  the  University  of  Kansas. 

All  communications  should  be  addressed  to 

The  Kansas  University  Science  Bulletin, 
Library  of  the  University  of  Kansas, 
Lawrence,  Kan. 


EDITORIAL  BOARD. 

S.  J..  Htjntbb,;  Chairman.  O.  O.  Stoland. 

R.  C.  MootiE,  Secretary.  W.  S.  Hunter. 

W.  C.  Stevens.  W.  J.  Baumgabtner. 

H.  C.  Teacy. 

Editorial  Note. — ^This  volume  was  assembled  and  published  under  the  edi- 
torial supervision  of  Dr.  H.  B.  Hungebford. 


APR  12  1924 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


L .  n. 


DEVOTED   TO 

THE  PUBLICATION  OF  THE  RESULTS  OF 

RESEARCH  BY  MEMBERS  OF  THE 

UNIVERSITY  OF  KANSAS 


Vol.  XIV 

ENTOMOLOGY  NUMBER  V. 


PUBLISHED  BY  THE  UNIVERSITY, 

LAWRENCE,  KANSAS. 

1922. 

9-4522 


CONTENTS  OF  VOLUME  XIV. 


No.  PaRo 

1.  Historical  Account  of  Department  of  Entomology 
(continued  from  Science  Bulletin  VIII).  H.  B. 
Hunger  ford   9 

2.  Withdrawn  from  publication. 

3.  The  Membracidae  of  Kansas  (Homoptera).  Plates 
I-VII.  P.  B.  Laivso7i 27 

4.  The  Genus  Acf)?op^e?'i4s  (Homoptera).  Plates  VIII- 
XII.     P.  B.  Laivson  Ill 

5.  The  Life  History  of  the  Toad  Bug  (Heteroptera). 
Plates  XIII-XIV.     H.  B.  Himgerford  143 

6.  A  New  Subterranean  Isopod  (Crustacea).  Plate 
XV.     H.  B.  Hjingerford  173 

7.  Studies  in  Cicadella  hieroghiphica  (Homoptera). 
Plates  XVI-XX.    Lucy  M.  Hackman 185 

8.  Ovipositors  of  Cicadellidse  (Homoptera).  Plates 
XXI-XXXIII.    P.  A.  Readio  213 

9.  Life  History  Notes  on  Two  Species  of  Saldidae  (Het- 
eroptera). Plates  XXXIV-XXXV.  Grace  OUve 
Wiley 299 

10.  A  Problem  in  the  Relation  of  Temperature  to  Rate 

of  Insect  Development.     P.  A.  Glenn  315 

11.  Some  Biological  Notes  on  Philippine  Zoology.  F.  X. 
Williams 327 

12.  Notes  on  Nesting  of  Polistea  (Hymenoptera). 
Divight   Isely    339 

13.  Five  New  Species  Belonging  to  Genus  HormoUta 
(Hymenoptera).  Plates  XXXVI-XXXVII.  W.  J. 
Phillips  and  Fred  W.  Poos 347 

14.  The  Urinary  System  of  Phlegethontius  sexta  (Lepi- 
doptera).    Plate  XXXVIII.     George  H.  Vansell  363 

15.  A  Brief  Resume  of  Investigations  Made  in  1913  on 
Trogoderma  inclusa  (Coleoptera).  Plates  XXXIX- 
XL.     A.  H.  Beyer  371 

16.  The  Larva  of  a  Chironomid  (Diptera).  Plates  XLI- 
XLIII.     P.  W.  Claassen  393 

(5) 


6  The  University  Science  Bulletin 

17.  Water  Insects  from  a  Portion  of  the  Southern  Utah 
Desert.    R.  C.  Moore  and  H.  B.  Himgerford  407 

18.  The  Nepidse  in  America  North  of  Mexico  (Heterop- 
tera) .    Plates  XLIV-LI.    H.  B.  Hungerford  423 

19.  A    Study   of   the   Relation    Between    Function   and 
Growth  in  Body  Cells.    Plates  LII.    M.  T.  Burrows    473 

20.  Notes    on    the    Biology    of    Curicta    (Heteroptera). 
Plate  LIII.     Mrs.  Grace  Olive  Wiley  505 

21.  Biology  and  Morphology  of  Lepyronia  quadrangu- 
laris    (Say) — Homoptera,   Cercopidse.     Plates   LIV 

to  LXII.     Kathleen  Doering  513 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  1— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  1.) 

ENTOMOLOGY  NUMBER  V 


CONTENTS: 

Historical  Account  of  Department  of  Entomology 
(continued  from  Science  Bulletin  VIII). 

H.  B.  Hungerford. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  1. 


Historical  Account  of  Department  of  Entomology. 

(Continued  from  Science  Bulletin  VIII), 

Brief  resume  of  the  work  of  the  Department  of  Entomology  of  the  University 
of  Kansas  during  the  past  quarter  centuiy. 

BY  H.  B.  HUNGERFORD. 

ENTOMOLOGICAL  work  has  been  in  progress  at  the  University 
of  Kansas  since  the  foundation  of  the  institution  in  1866.  The 
first  faculty  consisted  of  three  men,  one  of  whom  was  Dr.  Francis 
Huntington  Snow,  professor  of  mathematics  and  natural  science. 

Doctor  Snow,  while  widely  interested  in  birds  and  flowers,  gave 
early  evidence  of  a  special  fondness  for  the  study  of  insects. 
Through  his  efforts,  and  those  of  others  who  have  followed  him, 
there  has  been  established  what  is  perhaps  the  greatest  general 
collection  of  insects  to  be  found  in  connection  with  any  state  uni- 
versity in  America. 

In  1890  Doctor  Snow  became  chancellor  of  the  University,  and 
Dr.  Vernon  L.  Kellogg  was  appointed  to  the  entomology  work,  first 
as  assistant  professor  of  entomology,  and  later  as  associate  pro- 
fessor. 

Doctor  Kellogg  was  called  to  Stanford  University  in  1894,  and  for 
the  year  1895  the  entomological  work  was  in  charge  of  W.  A.  Snow, 
son  of  the  chancellor.       ^ 

In  1896  Prof.  S.  J.  Hunter  was  appointed  assistant  professor  of 
entomolog}^  and  placed  in  charge  of  the  department,  and  for  the 
past  quarter  of  a  century  he  has  directed  his  energies  towiird  the 
development  of  a  department  that  should  rank  among  the  strongest 
in  the  country.  During  this  span  of  years  the  department  has 
trained  many  students  and  grown  remarkably  in  its  material  equip- 
ment, both  through  the  greatly  enriched  entomological  collections 
and  its  mechanical  facilities  for  furthering  research  and  advancing 
instruction  in  entomology. 

(9) 


10  The  University  Science  Bulletin. 

The  teaching  staff  has  increased  from  one  to  five,  and  the  student 
roll,  from  a  few  to  more  than  250.  A  total  of  2,000  pages,  under 
276  titles,  have  been  published  by  members  of  the  department, 
and  twenty-three  scientific  expeditions  of  the  entomological  museum 
have  been  made  during  the  past  twenty-five  years.  These  accom- 
plishments, together  with  the  various  economic  state  problems  that 
have  arisen  from  time  to  time,  indicate  an  active  and  productive 
period  in  the  history  of  the  department. 

In  1914,  Science  Bulletin  issued  its  second  entomological  num- 
ber. This  was  dedicated  to  Doctor  Snow,  and  in  it  may  be  found 
a  chronological  account  of  the  activities  of  the  department  up  to 
and  including  the  year  1913.  As  a  matter  of  record,  additions  are 
made  below  to  the  various  sections  as  they  have  occurred  since 
that  year. 

CHRONOLOGICAL  REVIEW  OF  EVENTS  SINCE  1913. 

1913.  Mr.  George  Collett  appointed  Fellow  in  Entomology. 

F.  X.  Williams,  who  received  his  M.  A.  degree  in  this  department  in 
1913,  resigns  to  study  for  the  doctorate  at  Harvard. 

Professor  Hunter,  Assistant  Professor  Hungerford,  Mr.  George  Vansell 
and  Mr.  George  Collett  conduct  a  biological  survey  in  Wyoming, 
Utah  and  Montana  with  special  reference  to  distribution  and  biology 
of  grasshoppers  of  Kansas. 

1914.  P.  W.  Claassen  becomes  assistant  state  entomologist. 
Raymond  Beamer  becomes  assistant  curator  of  museum. 

Professor  Hunter,  Assistant  Professor  Hungerford,  Raymond  Beamer, 
Will  Brown  and  Fred  Poos  collect  along  the  Rio  Grande  in  south- 
western Colorado. 

1915.  Walter  Wellhouse  elected  fellow  in  entomology. 

Mr.  Raymond  Beamer,  B.  P.  Young,  Forrest  Anderson  and  Walter  Well- 
house  make  an  entomological  survey  of  the  counties  of  southeastern 
Kansas. 

B.  P.  Young  takes  charge  of  the  inseetary. 

1916.  Forrest  Anderson  elected  fellow  in  entomology. 
Professor  Hunter  takes  a  four  months'  lea^•e  of  absence. 

H.  B.  Hungerford  appointed  temporary  chairman  and  acting  state  en- 
tomologist. 

Hungerford,  Lawson  and  Wellhouse  go  to  Welhngton,  Kan.,  to  investi- 
gate green-bug  outbreak.  The  Federal  Bureau,  the  Kansas  Agricul- 
tural College  and  the  University  combine  to  study  the  green  bug 
and  methods  of  control.  Lawson  and  Wellhouse  represent  the  Uni- 
versity for  several  weeks  in  this  work. 

Mr.  Raymond  Beamer,  with  Grutzmacher,  Darby  and  Gardner,  make 
an  entomological  survey  of  some  southern  Kansas  counties. 

H.  B.  Hungerford  is  granted  a  year's  leave  of  absence  to  study  for  the 
doctorate  at  Cornell  LTniversity. 


Historical  Account.  11 

P.  W.  Claat;!?en  takes  Mr.  HungerfonlV  place  for  the  year. 
P.  B.  Lawson  becomes  assi-stant  instructor. 

1917.  H.  B.  Hungerford  inad(>  associate  professor  of  entomology. 
P.  B.  Lawson  made  instructor  in  entomology. 

Ruby  Hosford  takes  charge  of  inscctary. 

Dr.  C.  P.  Alexander  becomes  assistant  curator  of  entomological  museum. 

Miss  Eva  Batchman  appointed  assistant  instructor. 

William  Hoffmann  elected  fellow  in  entomology. 

Doctor  Ale.xander  conducts  a  party,  consisting  of  Harry  Fackler,  Wil- 
liam HotTmann  and  Scott  Johnson,  on  an  entomological  survey  of 
southwestern  Kansas. 

Canker-worm  work  in  cities  of  eastern  Kansas  conducted  by  Professor 
Hunter. 

1918.  p]ntire  University  reorganized  to  handle  S.  A.  T.  C.     Department  of 

entomology  teaches  three  sections  of  biology.    Miss  Weaverling  takes 
charge  of  insectary. 

1919.  William  Hoffmann  is  apiiointed  assistant  in  charge  of  insectary. 

P.  B.  Lawson  granted  degree  of  doctor  of  philosophy'  by  Kansas  LTni- 
versity;  major,  insect  taxonomy;  first  minor,  insect  morphology; 
second  minor,  systematic  botany.  Title  major  thesis,  "Cicadellidae 
of  Kansas." 

Miss  Itasca  Hilsman  elected  Fellow  in  Entomology. 

1920.  H.  B.  Hungerford  made  Professor  of  Entomology. 

P.  B.  Lawson  becomes  Assistant  Professor  of  Entomology. 
Mr.  William  Hoffmann  transferred  from  the  insectary  to  assistant  cura- 
tor of  museum. 
Mr.  Philip  A.  Readio  is  appointed  instructor  in  entomology. 
Miss  Lucy  Hackman  becomes  assistant  instructor. 
Miss  Itasca  Hilsman  reappointed  fellow  in  entomology. 

1921.  Mr.  Lawson  becomes  associate  professor  of  entomology. 
W.  J.  Brown  becomes  student  assistant  in  the  museum. 

W.  J.  Brown,  Jean  Linsdale  and  Robert  Guentert  continue  the  biological 

survey  in  northeastern  Kansas. 
P.  B.  Lawson  and  Raymond  Beamer  devote  some  time  to  the  study  of 

the  destructive  outbreak  of  pea  aphis  in  first-crop  alfalfa. 

1922.  Miss  Kathleen  Doering  appointed  fellow  in  entomology. 

P.  B.  Lawson  becomes  professor  of  entomology  and  assistant  dean  of 
the  College  of  Liberal  Arts  and  Science. 

Miss  Kathleen  Doering  appointed  scientific  illustrator. 

C.  Howard  Curran,  of  Orillia.  Canada,  appointed  research  fellow  in 
entomology. 

R.  H.  Beamer  reappointed  assistant  curator  of  the  entomological  col- 
lections. 


12  The  University  Science  Bulletin. 

ENTOMOLOGICAL  PAPERS  FROM  UNIVERSITY  OF  KANSAS. 

(Continued  from  Science  Bulletin  VIII.) 
ALEXANDER,  C.  P. 

No.      Date.  Title  of  paper. 

203.  1918 — Records  of  Japanese  Crane  Flies  (Diptera).    Am.  Ent.  Soc.  Amer., 

vol.  XI,  pp.  443-449. 

204.  1918 — New  Species  of  Tipuline  Crane  Flies  from  Eastern  Asia  (Tipulidse, 

Diptera).    Jour.  N.  Y.  Ent.  Soc.  vol.  XXVI,  pp.  66-75.- 

205.  1918 — A  New  Interpretation  of  the  Wing  Venation  of  the  Pedicine  Crane 

Flies  (Tipulidse,  Diptera).    Ent.  News,  vol.  XXIX,  pp.  201-205. 

206.  1918 — New  Species  of  Crane  Flies  from  California  (Diptera).    Ent.  News, 

vol.  XXIX,  pp.  285-288. 

207.  1918— New   Nearctic  Crane   Flies    (Tipulidse,  Diptera),  Part  IV.     Can. 

Ent.,  vol.  L,  pp.  60-71. 

208.  1918— New  Nearctic  Crane  Flies  (Tipulidse,  Diptera),  Part  V.    Can.  Ent., 

vol.  L,  pp.  158-165,  242-246. 

209.  1918— New  Nearctic  Crane  Flies    (Tipulidse,  Diptera),   Part  VI.     Can. 

Ent.,  vol.  L,  pp.  381-386,  411-416. 

210.  1919 — The  Crane   Flies   Collected  by   the   Canadian   Arctic  Expedition, 

1913-"18,  Report  of  the  Canadian  Arctic  E.xpedition,  1913-'18,  vol. 
Ill,  part  C,  pp.  IC  to  30C. 

211.  1919 — Notes  on  the  Genus  Dicranoptycha  Osten  Sacken.    Ent.  News,  vol. 

XXX,  pp.  19-22. 

212.  1919 — New  or  Little-known  Crane  Flies  from  Japan.  Part  I.     The  En- 

tomological Magazine,  Kyoto,  Japan,  vol.  Ill,  pp.  122-127. 

213.  1919— The  Biology  of  the  North  American  Crane  Flies  (Tipulidse.  Dip- 

tera),  Part  V.     Pomona   College  Jom-.  Ent.   and  Zool.,  vol.  XI. 
pp.  67-74. 

BEYER,   ADOLPH. 

214.  1922 — A  Brief  Resume  of  Investigations  Made  in   1913  on  Trogoderma 

inclusa.    Science  Bulletin,  vol.  XIV  (this  number). 

BEAMEB,  RAYMOND  H. 

215.  1916 — An    Easy    Method    of    Making    Insect   Labels.     Ent.    News,    vol. 

XXVIII,  p.  418. 

216.  1917— The  Oedipodinse  of  Kansas.    Bui.  of  the  Dept.'of  Ent.,  University 

of  Kansas,  No.  11.  pp.  51-126;  74  text  figures. 

CLAASSBN,  p.  \V. 

217.  1914 — Grasshopper  Control  in  the  Southern  Division   of  Kansas    (with 

Professor  Hunter).    Jour.  Ec.  Ent.,  vol.  VII,  No.  1,  pp.  73-81. 

218.  1917— The  Melanopli  of  Kansas.    Bui.  of  the  Dept.  of  Ent..  University 

of  Kansas,  No.  11,  pp.  5-50;  5  figures. 

CURRAN,  C.  HOWARD. 

219.  1922— On  the  Nemestrinid  Genus  Rhijncocephalus  (Nemestrinidse,  Dip- 

tera), Can.  Ent.,  vol.  LIV. 

220.  1922— Diptera  in  the  Collection  of  Miss  Frances  Long   (supplement  to 

paper  by  Miss  Frances  Long  on  Plant  Pollination  in  Insects) .    (In 
press.) 


Historical  AccorxT.  13 

DOERINC;.    KATHLEEN. 

221.  1922 — Biology  and  Morpholojiy  of  Lcpi/rouia  qundrangularis  (Say).    Sci- 

ence Bullf^fiii.  \(il.  Xn'  (this  number). 

EMERY,    \V.   T. 

222.  1914 — Morphologj-  and  Biologj'  of  Simulium  vittatum  and  its  Distribu- 

tion in  Kansas.    Science  Bulletin  VIII;  15  plates. 

H.ACKMAN,  LUCTi'. 

223.  1922 — Studies    in    Cicadella   hieroglyphica.     Science    Bulletin,   vol.  XIV 

(this  number) ;  5  plates. 

HUNGERFOKD,   H.  B. 

224.  1914 — Anatomy  of  Simulium  vittatum.     Science  Bulletin,  vol.  VIII,  pp. 

365-382;  3  plates. 

225.  1914 — Notes  on  Coleoptera  from  Western  Kansas  (with  F.  X.  Wilhams). 

Ent.  News,  vol.  XXV,  pp.  1-9;  2  plates, 

226.  1915— A  Parasite  of  Cottonwood ■  Borer  Beetle.     Ent.  News,  vol.  XXVI, 

p.  135. 

227.  1916 — Sciara  Maggots  Injurious  to  Potted  Plants.     Jour.  Ec.  Ent.,  vol. 

IX,  pp.  538-549;  2  plates, 

228.  1917 — Brief    Laboratory    Outline    for    Introductory    Entomology.     State 

Printer;  18  pages. 

229.  1917 — Egg-laying    Habits    of    a    Back    Swimmer,    Buenoa    margaritacea 

Bueno.    Ent.  News,  vol.  XXVIII,  pp.  174-183;  1  plate. 

230.  1917 — Notes  Concerning   Food   Supply  of   Some   Water  Bugs.     Science 

N.  S.,  XLV,  pp.  336-337. 

231.  1917— Food  Habits  of  Corixids.    Jour.  N.  Y.  Ent.  Soc,  vol.  XXV,  pp.  1-5; 

1  plate. 

232.  1917 — The  Life  History  of  a  Back  Swimmer,  Notonccta  undulata  Say. 

Ent.  News,  vol.  XXVIII,  pp.  267-278,  2  plates. 

233.  1917 — The   Life    Histoiy    of   Mesovelia    mulsanti    White.      Psyche,    vol. 

XXIV,  pp.  73-84;  1  plate. 

234.  1917— The  Life  History  of  a  Boatman.    Jour.  N.  Y.  Ent.  Soc,  vol.  XXV, 

pp.  112-122;  1  plate. 

235.  1918 — Notes  on  the  Ovii)osition  of  Some  Semiaquatic  Hemiptera,     Jour, 

N,  Y.  Ent.  Soc,  vol,  XXVI,  pp,  12-18;  1  plate. 

236.  1918 — Concerning  the   Oviposition  of  the   Notonecta.     Ent,   News,   vol. 

XIX,  pp.  241-243;   1  plate, 

237.  1919 — Biology    and    Ecology    of    Aquatic    and    Semiaquatic    Hemiptera. 

Science  Bulletin,  vol.  XI,  pp.  3-328,  33  plates, 

238.  1919 — Male  Genitalia  as  Characters  of  Specific  Value  in  Certain  Crypto- 

cerata.    Science  Bulletin,  vol.  XI,  pp,  329-332;  2  plates. 

239.  1919 — Biological  Notes  on  Tetradonema  plicans  Cobb,  a  Nematode  Para- 

site of  Sciara  coprophila  Lint,     Jour,  of  Parasitology,  vol.  V,  pp. 
176-192;  1  plate,  3  text  figures. 

240.  1919 — Tables  for  Determining  Types  and  Breeds  of  Domesticated  Ani- 

mals (third  edition).    Comstock  Publishing  Co.,  Ithaca,  N.  Y.;  38 
pp.,  3  plates. 


14  The  University  Science  Bulletin. 

241.  1920 — Laboratory  Outline  for  Course  in  Introductorj^  Entomology    (re- 

vised and  enlarged).    World  Publishing  Co.;  39  pp. 

242.  1922 — Life  History  of  the  Toad  Bug,  Gelastocorus  oculatns  Fabr.    Science 

Bulletin,  vol.  XIV;  2  plates  (this  number). 

243.  1922 — A  new  Subterranean  Isopod  from  Kansas.     Science  Bulletin,  vol. 

XIV;  1  plate  (this  number). 

244.  1922 — A  Review  of  the  Past  Quarter  Century  of  Entomology  in  Kansas 

University.    Science  Bulletin,  vol.  XIV  (this  number). 

245.  1922— The  Nepidae  of  North  America.     Science  Bulletin,  vol.  XIV;    8 

plates  (this  number). 

246.  1922 — Oxyhsemoglobin  Present  in  the  Back  Swimmer,  Buenoa  margari- 

tacea  Bueno.    Can.  Ent.,  vol.  LIV. 

247.  1922 — Saldoidea  slossoni,  new  var.  voileyii.    Bui.  Brooklyn  Ent.  Soc,  vol. 

XVII,  page  64  (Apr.). 

248.  1922 — Notable  Additions  to  Entomological  Library  at  Kansas  University. 

Bui.  Brooklyn  Ent.  Soc.     (Accepted  for  publication.) 

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254.  1914 — Control  Measures  for  the  Native  Grasshoppers  and  Chinch  Bugs. 

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257.  1915 — Some  Economic  Results  of  the  Year.     Jour.  Ec.  Ent.,  vol.  VIII, 

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260.  1917 — Practical   Insecticides   and   Proper  Application.     Trans.   Kan.   St. 

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261.  1917 — Spring  Cankerworm:    An  Orchard  and  City  Problem.    Trans.  Kan. 

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262.  1918 — Municipal  Control  of  Spring  Cankerworm.    Jour.  Ec  Ent.,  vol.  XI, 

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263.  1918— The  Wood  Lot.    Bien.  Rep.  Kan.  St.  Hort.  Soc,  vol.  XXXV,  pp. 

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HisTOHK  AL  Account.  15 

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271.  1917— Dragon  Flies  of  Kansas.     Bui.  of  the  Dept.  of  Ent.,  University  of 

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L.UVSON,    P.    B. 

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275.  1922 — List  of  the  Cicadellidae  of  Kansas  (Homoptera).    Trans.  Kan.  Acad. 

Sci..  vol.  XXX,  pp.  331-336. 

276.  1922 — List   of  the  Grasses  of  Douglas  County.     Trans.  Kan.  Acad.  Sci., 

vol.  XXX,  pp.  336-339. 

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278.  1922 — The    Membracidse    of    Kansas.     Science    Bulletin,    vol.   XIV    (this. 

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Insecticide.    Jour.  Ec.  Ent.,  vol.  IX,  No.  1,  pp.  169-170. 

280.  1916 — Formulas    for    Destroying    Injurious    Insects    and    Plant    Diseases.. 

University  of  Kansas  Circular,  No.  4. 

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WILEY,   GR.ACE   OLIVE. 

282.  1922— Life  History  Notes  on  Two  Species  of  Saldida>.     Science  Bulletin, 

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Can.  Ent.,  vol.  L,  No.  8,  pp.  267-277;  figures,  2. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  3— October,  1922. 

(Whole  Series,  Vol.  XXIV,   No.    3.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

The  Membr.'Vcid.e  of  K.ans.as  (Homopter.a) P.  B.  Lawson. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


TABLE  OF  CONTENTS. 

PAGE 

Introduction 31 

Distribution 32 

Structural  characteristics 36 

The  male  genitalia 38 

Phylogeny  of  the  family 41 

Life  history 41 

Economic  importance 43 

List  of  Kansas  species 44 

Systematic  treatment  of  Kansas  species 45 

Subfamily  Centrotin.e 45 

The  genus  Microcentras 45 

Subfamily  Membracin^ ^ 47 

The  genus  Campylenchia 47 

The  genus  Enchenopa 48 

Subfamily  Smiliin.e 50 

The  genus  Ceresa 51 

The  genus  Stictocephala 59 

The  genus  Acutalis 62 

The  genus  Micrutalis 63 

The  genus  Carynota 64 

The  genus  Thelia 65 

The  genus  Glossonotus 67 

The  genus  Heliria 68 

The  genus  Telamona 69 

The  genus  Telamonanthc 77 

The  genus  Archasia 78 

The  genus  Smilia 79 

The  genus  Cyrtolobus 80 

The  genus  Ophiderma 87 

The  genus  Vanduzea 89 

The  genus  Entylia 90 

The  genus  Publilia 91 

Index 107 

(29) 


THE  KANSAS  UNIYEBSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  3. 


The  Membracidae  of  Kansas. 

By  P.   B.   LAWSON, 

Professor  of  Eiitoniologj',  University  of  Kansas. 

INTRODUCTION. 

THE  writer  has  been  interested  in  recent  years  in  a  systematic 
study  of  the  Homoptera  of  Kansas.  He  has  previously  pub- 
hshed  papers  on  the  Coccidse,  Cicadelhda^  and  Cicadidae  of  the  state, 
listing  over  300  species  of  the  members  of  these  families  known  to 
occur  within  the  borders  of  Kansas. 

The  membracids  of  the  state  have  been  previously  studied  by 
Miss  Hazel  Branch,  who  in  1914  published  a  paper  in  the  Kansas 
University  Science  Bulletin,  volume  8,  on  the  biology  of  the 
Membracidse  of  Kansas.  In  that  paper  she  listed  nineteen  species 
from  the  state.  Since  that  time  quite  a  little  collecting  has  been 
done,  until  to-day,  including  some  species  taken  around  Kansas 
City,  Mo.,  which  species  are  therefore  sure  to  occur  in  eastern  Kan- 
sas also,  there  are  records  of  the  occurrence  of  at  least  fifty-five 
species  of  tree  hoppers  in  our  fauna.  Further  collecting  will  of 
course  reveal  a  goodly  number  of  additional  species,  but  the  writer 
has  thought  it  advisable  to  bring  our  data  up  to  date  at  this  time. 

The  writer  is  greatly  indebted  to  Dr.  W.  D.  Funkhouser,  of  the 
University  of  Kentucky,  who  has  made  many  determinations  for 
him  and  been  very  generous  in  giving  helpful  suggestions  and  en- 
couragement during  the  course  of  this  study.  Moreover,  his  paper 
on  the  biology  of  the  Membracidse  of  the  Cayuga  Lake  Basin  has 
been  very  freely  drawn  upon,  especially  in  the  use  of  the  technical 
descriptions. 

Through  the  kindness  of  Professors  Geo.  A.  Dean  and  Roger 
Smith,  the  records  of  the  Kansas  State  Agricultural  College  are  in- 
cluded in  this  paper,  most  of  the  records  from  Riley  county  and  sev- 
eral others  being  from  that  collection. 

rsi) 


32  The  University  Science  Bulletin. 


DISTRIBUTION. 

The  Membracidse,  though  primarily  a  tropical  and  subtropical 
family,  are  nevertheless  found  widely  scattered  and  are  well  repre- 
sented in  temperate  regions.  Authorities  are  agreed  that  they  are 
best  represented  in  the  fauna  of  Central  and  South  America,  but  it 
seems  certain  that  when  the  fauna  of  Africa  and  Southern  Asia  have 
been  as  carefully  studied  that  these  regions  will  also  be  found  to  be 
very  rich  in  these  insects. 

Dr.  W.  D.  Funkhouser  gives  the  following  distribution  of  the  fam- 
ily according  to  geographical  life  zones: 

Palearctic  region: 

(Europe,  the  temperate  parts  of  Asia,  and  the  north  of  Africa;  Iceland  and 
the  ishinds  of  the  Atlantic;  limited  by  the  Himalaj-as.) 

Very  poorly  represented.  Only  two  or  three  genera  on  the  entire  continent 
of  Europe,  but  two  species  in  Great  Britain,  two  species  in  Russia,  and  none 
reported  from  Iceland.  A  few  in  northern  Africa,  chiefly  forms  that  have 
migrated  from  the  south. 

Ethiopian  region: 

(Africa  and  its  islands,  except  the  northern  parts;  Arabia.) 
Rich  in  genera  and  species.    Little  work  has  been  done  on  these  forms  of 
the  family,  but  there  is  evidence  of  an  abundant  membracid  fauna. 

Oriental  region: 

(India  and  the  East  Indies.) 

Extremely  rich  both  in  number  of  forms  represented  and  in  number  of  in- 
dividuals.   The  center  of  distribution  for  the  subfamily  Centrotinae. 

Australian  region: 

(Australia,  New  Zealand  and  neighboring  islands.) 

Well  represented  by  rather  distinct  forms.  The  region  has  been  fairly  well 
worked  and  has  yielded  a  large  number  of  species. 

Nearctic  region: 

(America  north  of  Mexico;  Greenland). 

Forty  or  fifty  genera,  gradually  becoming  less  abundant  northward.  A 
few  species  common  in  Canada  as  far  north  as  Peny  Sound.  None  reported 
from  Greenland. 

Neotropical  region: 

(Mexico,  West  Indies,  Central  and  South  America.) 

The  most  important  of  all  the  regions  for  the  Membracidse.  Central 
America  and  the  northern  part  of  South  America  have  yielded  as  many  spe- 
cies as  all  the  rest  of  the  world  together. 

Altogether  over  300  genera  have  been  erected,  which  contain 
something  like  1,500  species.     Many  of  these  genera  are  found  in 


Lawson:    Membracid.e  of  Kansas.  33 

more  than  one  of  I  he  abox'c  regions,  but  the  number  of  !>ueh  species 
is  seemingly  rather  limited.  Thus  the  American  genera  Ceresa  and 
Stictocephala,  while  well  represented  in  both  the  Nearctic  and  Neo- 
tropical regions,  scarcely  have  a  single  species  that  occurs  in  both 
regions.  On  the  other  hand,  it  is  well  known  that  some  forms,  such 
as  Micndalis  calva  (Say),  are  found  in  both  the  United  States  and 
the  West  Indies. 

In  North  America  the  members  of  this  family  are  best  represented 
in  Mexico.  As  we  advance  northward  they  become  fewer  and  fewer 
till  comparatively  few  species  are  found  in  Canada,  where  they 
reach  their  northern  limit.  Van  Duzee  lists  41  genera  in  his  cata- 
logue of  the  Hemiptera  -of  North  America  north  of  Mexico.  It  is 
perhaps  safe  to  say  that  the  number  of  genera  for  this  region  does 
not  exceed  fifty.  The  number  of  species  is  listed  at  185;  which  com- 
pares very  favorably  with  the  25  cercopids  and  the  74  cicadas,  but 
falls  far  short  of  the  357  fulgorids  and  the  698  cicadellids  which  oc- 
cur in  the  same  territory.  Thus  it  is  seen  that  in  the  United  States 
this  family  stands  midway  in  membership  among  the  five  families 
of  the  Homoptera-Aucheinorhynchi. 

So  far  the  membracid  fauna  of  Kansas  is  known  to  be  repre- 
sented by  20  genera.  Further  collecting  will  doubtless  reveal  the 
presence  of  several  other  genera,  and  of  course  the  number  of  spe- 
cies will  also  be  increased,  for,  after  all,  there  has  been  rather  little 
attention  paid  to  this  family  in  the  work  of  past  collectors  in  this 
state.  This  is  shown  by  the  fact  that  only  about  50  out  of  the  105 
counties  of  the  state  have  as  yet  furnished  us  with  specimens.  How- 
ever, these  counties  are  so  well  distributed  as  to  give  us  a  very  fair 
idea  of  the  membracid  fauna  of  the  entire  state. 

The  genera  Telamona  and  Cyrtolobus  show  the  most  species  in 
the  state,  each  having  some  ten  or  eleven  species.  These  genera  are, 
however,  not  the  ones  that  are  most  commonly  seen,  for  both 
Stictocephala  and  Ceresa  are  far  more  abundant  as  to  individuals, 
Ceresa  bubalus  (Fabr.)  being  by  far  the  commonest  species  in  the 
state.  Close  to  the  two  latter  genera  comes  Micrutalis,  the  species 
calva  having  been  taken  in  about  fifteen  counties.  Campy lenchia 
latipes  is  also  very  common  throughout  the  alfalfa  fields  of  the 
state.  Vanduzea  triguttata  (Burm.)  occurs  in  enormous  numbers 
when  taken,  as  does  Entylia  concisa  (Walk.),  but  hitherto  these 
species  have  not  been  taken  in  many  places  within  our  borders. 


34  The  University  Science  Bulletin. 


EXPLANATION  OF  PLATE  I. 

1.  Pronotum  and  head  of  Ceresa  bubalus.  sp,  suprahuraerals ;  m,  meto- 
pidium;  ha,  humeral  angles. 

2.  Cephalic  aspect  of  head  of  Ceresa  bubahis.  es,  epicranial  suture;  o, 
ocellus;  v,  vertex;  e,  compound  eye;  g,  gena;  lo,  lora;  cl,  clypeus;  I,  labrum; 
ep,  epipharynx;  la,  labium. 

3.  Caudal  aspect  of  head  of  Cerem  bubalus.  oc,  occiput;  of,  occipital 
foramen;  pg,  postgena;  c,  compound  eye;  la,  labium. 

4.  Lateral  aspect  of  head  of  Ceresa  bubalus.  oc,  occiput;  v,  vertex;  e, 
compound  eye;  g,  gena;  cl,  clypeus;  pg,  postgena;  I,  labrum;  ep,  epipharynx; 
mx,  maxillary  stylets;  md,  mandibular  stylets;  la,  labium. 

5.  Pronotum  of  Ceresa  bubahis.  sp,  suprahumerals;  ha,  humeral  angles; 
m,  metopidium;  dc,  dorsal  carina;  pp,  posterior  process. 

6.  Metathoracic  leg  of  Ceresa  bubalujs.  c,  coxa;  tr,  trochanter;  /,  femur; 
t,  tibia;  ta,  tarsus. 

7.  Prothoracic  leg  of  Ceresa  bubalus. 

8.  Prothoracic  leg  of  Campylenchia  latipes. 

9.  Tegmen  of  Ceresa  bubalus. 
10.   Hind  wing  of  Ceresa  bubalus. 

IL   Ventral  aspect  of  abdomen  of  male  Ceresa  bubalus.    2-9,  sternites;  2-8, 
pleurites;  py,  p.ygofer  or  ninth  tergite;  Iv,  lateral  valve. 

12.  Ventral  aspect  of  abdomen  of  female   Ceresa  bubalixs.     2-7,  sternites; 
2-S,  pleurites;  pif,  pygofer  or  ninth  tergite;  ovi,  ovipositor. 

13.  Lateral  aspect  of  abdomen  of  male  Ceresa  bubalus.     1-11,  tergites;  2-9, 
sternites;  Iv,  lateral. valve. 

14.  Lateral  aspect  of  abdomen  of  female  Ceresa  bubalus.    1-9,  tergites;  2-7, 
sternites. 


Lawson:    ISIembracid.e  of  Kansas. 


35 


PLATE  I. 


36  The  University  Science  Bulletin. 


STRUCTURAL  CHARACTERISTICS. 

The  outstanding  characteristic  of  the  membracids  is  their  re- 
markably developed  pronotum.  This  sclerite  is  usually  enlarged 
so  as  to  cover  most  of  the  thorax  and  much  of  the  abdomen,  and 
while  in  our  species  it  is  often  extended  into  rather  regularly  formed 
processes,  yet  in  many  tropical  species  these  processes  assume  the 
most  irregular  and  grotesque  shapes.  In  the  subfamily  Centrotinse, 
however,  the  pronotum  is  not  thus  enlarged,  but  is  so  small  that  the 
greater  part  of  the  scutellum  is  left  exposed. 

As  in  all  insects,  the  body  is  divided  into  three  regions.  The  head 
is  quite  characteristically  homopterous,  fitting  against  the  coxae  of 
the  prothoracic  legs,  between  which  lies  the  beak.  At  its  lateral 
extremities  are  the  well-developed  compound  eyes,  while  the  simple 
eyes  or  ocelli  are  found  on  the  i  ertex,  the  paired  sclerite  forming 
the  greater  part  of  the  cephalic  aspect  of,  the  head.  The  ocelli  are 
always  paired  and  in  a  line  with  each  other  and  the  compound  eyes, 
though  their  distance  from  each  other  and  from  the  compound  eyes 
varies. 

The  setaceous  antennce  are  found  under  the  margin  of  the  vertex 
on  either  side  of  the  clypeus.  The  first  three  segments  are  large,  but 
the  rest  of  the  organ  is  composed  of  a  large  number  of  small 
segments  which  decrease  in  size  toward  the  apex. 

The  vertex  is  a  paired  sclerite  lying  on  either  side  of  the  base  of 
the  Y-shaped  epicranial  suture.  The  presence  of  this  suture  would 
indicate  a  mlore  primitive  position  for  this  family  than  for  the 
Cicadellidae,  where  no  signs  of  it  are  present. 

Between  the  arms  of  the  epicranial  suture  and  forming  the  apex 
of  the  head  is  the  clypeus.  This  sclerite  is  often  characteristic  in 
the  length  of  its  apex  and  in  its  relation  to  the  lateral  margins  of 
the  vertex.  At  its  apex  it  is  turned  backward,  forming  a  rounded 
lobe,  caudad  of  which  are  the  genoi.  The  latter  sclerites  are  seen 
to  form  the  lateral  portions  of  the  head,  extending  between  the  eyes 
and  the  clypeus,  and  bearing  the  antennae.  The  postgence  occupy 
most  of  the  caudal  aspect  of  the  head,  forming,  with  the  dorsal 
occipital  sclerites  or  occiput,  most  of  the  boundary  of  the  occipital 
foramen.  The  lorce  are  found  on  either  side  of  the  clypeus  and 
labrum  as  two  more  or  less  distinct  sclerites  with  rounded  lateral 
margins. 

The  beak  or  labium  consists  of  three  segments,  as  in  other 
Homoptera,  and  contains  in  a  groo^'e  the  mandibular  and  maxillary 


Lawson:    Membracid^  of  Kansas.  37 

stylets.  The  labrum  is  large  aiul  cauchul  of  the  elypeus,  bearing  the 
more  or  less  membranous  epipharynx  at  its  tip.  All  the  mouth 
parts  are  typical  of  the  ordinary  sucking  phytophagous  insect. 

The  thorax,  as  in  all  insects,  consists  of  three  segments.  For  our 
purposes  only  a  brief  discussion  of  the  dorsal  sclerite  of  the  pro- 
thorax  will  be  necessary.  This  sclerite,  the  pronotum,  as  already 
mentioned,  is  the  chief  characteristic  of  the  family.  It  is  one  of 
the  finest  examples  of  the  biological  phenomenon  of  orthogenesis 
that  is  to  be  found.  In  our  North  American  membracids  it  is  com- 
paratively simple  as  compared  with  its  appearance  in  many  Neo- 
tropical genera,  such  as  Sphongophorus  and  Cyphonia.  But  even  in 
our  fauna  it  assumes  such  diverse  forms  as  to  give  us  the  best 
generic,  and  often  also  specific  characters  for  the  classification  of  the 
members  of  this  family.  The  following  parts  have  been  named  and 
are  used  in  the  descriptions  which  follow: 

1.  Metopidium.  The  cephalic  area  ,of  the  pronotum,  extending 
from  the  base  of  the  head  to  the  front  of  the  dorsum. 

2.  Humeral  angles.  The  angles  of  the  pronotum  just  above  the 
bases  of  the  wings. 

3.  Suprahumerals.  The  lateral  projections  above  the  humeral 
angles. 

4.  Dorsal  carina.  The  ridge  usually  extending  the  length  of  the 
pronotum. 

5.  Posterior  process.  The  caudally  directed  portion  of  the 
pronotum. 

The  three  segments  of  the  thorax  each  bear  a  pair  of  legs,  which 
are  composed  of  the  usual  segments,  the  tarsi  being  three-segmented. 
The  coxce  are  usually  stout,  the  trochanters  rather  small  and  bent, 
the  femora  large  and  stout,  and  the  tibice  usually  long,  slender  and 
hairy.  In  the  subfamily  Membracinae  and  in  some  Centrotinae,  the 
tibiae  are  wide  and  flat  or  foliaceous,  especially  so  in  the  prothoracic 
and  mesothoracic  legs. 

The  last  two  thoracic  segments  each  bear  a  pair  of  wings,  which 
are  of  value  in  sj^steraatic  work,  not  only  in  their  venation,  but  also 
in  their  relation  to  the  pronotum. 

The  abdomen  consists  of  eleven  segments,  the  last  two  forming 
the  anal  tube.  Most  of  these  segments  show  a  distinct  tergite, 
pleurites  and  sternite.  Of  these  sclerites  the  tergite  is  by  far  the 
largest,  forming  both  the  dorsal  and  lateral  portions  of  the  seg- 
ment; and  in  the  case  of  the  female  ninth  segment,  nearly  inclosing 
the  segment,  coming  clear  down  to  the  sides  of  the  ovipositor,  in 


38  The  University  Science  Bulletin. 

this  sex  there  showing  externally  but  six  sternites,  those  of  the 
second  to  seventh  segments,  inclusive.  In  the  male  the  ninth  tergite 
is  not  as  large,  for  in  this  sex  the  sternites  of  segments  two  to  nine, 
inclusive,  all  show  clearly.  This  tergite  bears,  however,  on  either 
side,  a  pair  of  broad  lobes  or  plates  which  Funkhouser  calls  the 
lateral  valves.  These,  he  thinks,  may  be  the  pleura  of  the  ninth 
segment  and  of  use  in  copulation.  They  may  project  caudad  or 
turn  mesad — a  character  of  generic  value.  They  frequently  bear 
variously  shaped  processes  or  teeth,  which,  as  far  as  the  writer  has 
observed,  are  constant  in  form  and  position  for  the  various  species, 
and  are  therefore  good  specific  characters. 

The  sternite  of  the  seventh  segment  of  the  female  differs  in  differ- 
ent species,  and  is  therefore  of  systematic  value,  as  is  also  the 
ninth  sternite  or  sternal  plate  of  the  male.  The  latter  is  constant 
in  shape  and  extent  of  apical  splitting  within  the  species,  and  this 
makes  it  of  value  as  a  taxonomic  character. 

THE  MALE  GENITALIA. 

As  in  the  Cicadellidae,  the  abdomen  of  the  male  ends  in  what 
Sharp  calls  the  "terminal  chamber"  in  the  case  of  the  Pentatomidse. 
This  chamber  is  bounded  above  by  the  anal  tube  and  ninth  tergite, 
laterally  by  the  lateral  valves,  and  ventrally  by  the  ninth  sternum. 
Within  this  chamber  are  found  the  genital  organs  of  the  male,  their 
position  and  structure  being  exactly  homologous  with  the  male 
genitalia  of  the  leaf  hoppers. 

The  styles  or  claspers  are  always  paired,  both  members  of  the  pair 
being  alike,  and  fastened  to  the  sternal  plate  by  apparently  passing, 
near  their  middle,  through  the  membrane  forming  the  dorsal  surface 
of  the  plate.  The  cephalic  portion  of  the  clasper  projects  into  the 
abdominal  cavity,  frequently  reaching  the  sixth  segment,  though 
usually  extending  only  as  far  as  the  eighth  or  ninth.  It  does  not 
seem  to  vary  much  in  the  different  species.  The  apical  portion, 
however,  is  characteristic  in  its  shape  and  apical  structure,  some- 
times being  nearly  straight,  at  other  times  more  or  less  strongly 
curved,  and  ending  either  in  a  plain  acute  point  or  in  variously 
shaped  and  toothed  extremities.  So  far  the  writer  has  found  no  two 
species  in  w'hich  the  styles  are  alike  in  both  size  and  structure. 

Canon  Fowler,  in  the  "Biologia  Centrali-Americana,"  states  that 
in  the  genus  Ceresa  the  styles  are  long  and  pointed,  while  in  the 
genus  Stictocephala  they  are  short  and  obtuse  at  the  apex.  This 
is  undoubtedly  frequently  true,  but  the  writer  has  found  that  it  does 


Lawson:    Membracid.e  of  Kansas.  39 

not  always  hold  in  tlu'  members  of  these  genera  as  the  latter  are  now 
constituted.  Thus,  while  in  Stictocephala  festina  the  above-men- 
tioned condition  holds  good,  it  is  not  true  of  S.  lutea,  for  the  latter 
possesses  styles  just  as  long  and  pointed  as  in  several  members  of 
Ceresa,  notably  C.  bubalus. 

As  in  the  Cicadellida>,  the  stvles  are  united  bv  a  small  sclerite, 
whicii  \-aries  in  shape  in  the  various  genera  and  species,  but  is 
always  of  the  same  shape  within  the  species.  This  sclerite  the 
writer  has  called  the  "style-oedagus  connective,"  or  more  simply, 
the  "connective.''  The  former  name  shows  its  usual  function  in  the 
leaf  hoppers,  for  usually  it  unites  not  only  the  two  styles,  but  at  its 
distal  end  is  united  to  the  base  of  the  cedagus.  This  condition  pre- 
vails in  all  the  tree  hoppers  thus  far  examined.  Its  relation  to  the 
styles,  however,  is  apparently  different  in  the  two  families,  for  in 
the  leaf  hoppers  each  style  sends  out  a  distinct  chitinous  process  to 
which  it  is  fastened,  while  in  the  tree  hoppers  this  chitinous  process 
is  not  usually  present,  but  rather  each  style,  at  the  point  of  attach- 
ment to  the  connective,  gives  off  a  dorsal  membranous  fold  which 
unites  with  the  connective,  the  two  folds  appearing  heart-shaped 
basally  when  viewed  from  above. 

In  all  the  species  so  far  examined  the  connective  appears  as  a 
rather  thin,  usually  fiat  sclerite,  frequently  showing  a  distinct  tend- 
ency to  fold  longitudinally  along  a  median  keel,  thus  drawing  the 
two  styles  closer  together.  It  is  usually  quite  small,  but  in  some 
forms,  notably  Ceresa  borealis,  it  is  well  developed. 

The  writer  has  not  tried  to  determine  the  morphological  status  of 
the  connective.  Kornhauser  calls  it  the  ninth  sternite.  Funkhouser 
does  not  mention  it,  but  feels  that  the  last  sternal  plate  or  the  valves 
are  the  ninth  sternite.  That  it  is  a  sternite  cannot  be  doubted,  and 
the  writer  hopes  to  discuss  its  exact  position  in  a  later  paper  on  the 
genitalia  of  the  Homoptera-Auchenorhynchi. 

The  cedagus,  or  perns  sheath,  seen  laterally,  is  usually  a  V-shaped 
organ,  the  mouth  of  the  V  looking  caudad  or  dorsad.  The  anterior 
or  dorsal  arm  extends  up  to  the  base  of  the  anal  tube,  the  latter 
always  being  fastened  to  it,  usually  at  a  slight  excision  near  its 
apex.  As  far  as  our  present  studies  go,  the  shape  of  this  arm  seems 
to  vary  more  or  less  even  wathin  the  species,  sometimes  being  rather 
narrow  and  straight,  then  changing  till  it  is  frequently  quite  broad 
and  variously  bent.  The  lower  or  posterior  arm  is  the  penis  sheath 
proper.  The  penis  enters  it  just  above  the  usual  basal  angle  by 
which  the  cedagus  is  united  to  the  connective.    This  basal  angle  may 


40  The  University  Science  Bulletin. 

often  be  prolonged  into  a  distinct  cephalic  process.  This  ventral  arm 
varies  greatly  in  different  genera  and  species,  but,  as  far  as  our 
observations  go,  is  always  constant  in  shape  within  the  species.  It 
usually  extends  more  or  less  dorso-caudad,  though  sometimes  its 
apex  is  bent  distinctly  dorsad  and  may  even  extend  dorso-cephalad. 
The  course  of  the  penis  through  this  arm  is  usually  easy  to  trace  in 
caustic  potash  specimens. 

In  the  genera  Ceresa  and  Stictocephala  the  functional  orifice  is 
usually  along  the  ventral  side  of  this  arm  and  at  quite  a  distance 
from  the  apex.  This  opening  in  these  genera  is  usually  quite  long 
and  is  guarded  by  a  distinct  membranous  external  sheath.  In  S: 
jestina,  however,  the  penis  opens  apically,  while  in  Campylenchia, 
Telamona  and  others  it  opens  at  or  near  the  apex.  In  every  case, 
however,  the  opening  seems  to  be  guarded  by  the  external  membrane 
and  is  constant  in  position  within  the  species.  The  shape  of  the 
apex  also  is  constant  for  the  species ;  in  fact,  the  whole  organ  seems 
to  afford  an  excellent  specific  character  for  use  in  systematic  work. 
Canon  Fowler's  characterization  of  the  cedagus  of  Stictocephala 
does  not  hold  good  for  all  the  species  of  the  genus. 

The  cephalic  portion  of  the  styles  and  the  dorsal  arm  of  the 
cedagus  are  practically  internal  parts.  It  seems  as  though  these 
parts  have  distinct  layers  added  to  them  at  various  intervals,  thus 
adding  to  the  length  of  the  former  and  to  the  width  of  the  latter 
This  may  account  for  the  variable  condition  seen  in  these  organs, 
even  within  the  species.  The  terminal  portions,  however,  seem  to 
be  constant. 

The  sternal  plate,  both  as  to  shape  and  the  extent  of  its  division 
into  two  parts,  is  an  excellent  and  readily  available  character  for 
taxonomic  work.  It  may  be  the  ninth  sternite,  but  the  question  as  to 
its  possible  homology  with  abdominal  appendages  at  once  arises. 
At  any  rate,  it  seems  to  be  exactly  homologous  with  the  plates  of  the 
Cicadellidse. 

Further  work  with  these  genital  organs  will  undoubtedly  lead  to 
their  greater  use  in  systematic  work.  The  writer  has  found  that 
specimens  identified  as  identical  can  frequently  be  separated  and 
properly  classified  by  a  study  of  these  organs.  It  is  to  be  hoped  that 
the  various  genera  will  be  studied  from  this  standpoint  in  the  days 
to  come.  If  they  are  the  writer  feels  that  several  generic  changes 
will  be  made.  Thus  it  would  seem  that  the  genus  Stictocephala 
should  be  divided,  for  the  genitalia  of  S.  festina  and  S.  lutea  cannot 
possibly  belong  to  members  of  the  same  genus.    On  the  other  hand, 


Lawson:    Membraoid.t:  of  Kansas. 


41 


the  genitalia  of  the  genera  Tclamona  and  Tclonaca  are  so  near  alike 
that  it  seems  certain  that  the  latter  should  never  have  been  separated 
from  the  formei-. 

PHYLOGENY  OF  THE  FAMILY. 

The  membracids  belong  to  that  group  of  the  Homoptera  known 
as  the  Auchenorhynchi,  which  differs  from  the  rest  of  the  members 
of  the  order,  the  Sternorhynchi,  in  that  in  the  latter  the  mouth  parts 
seeminglj'-  arise  from  between  the  prothoracic  legs,  instead  of  from 
the  head,  as  in  the  former. 

In  this  group  are  five  families,  arising  from  three  different  stems, 
of  which  it  seems  clear  that  the  Cicadidae  arise  from  the  lowest  stem 
and  the  Fulgoridse  from  the  highest.  In  between  these  two  families 
are  the  closely  related  membracids,  cicadellids  and  cercopids.  As 
pointed  out  in  his  paper  on  the  Cicadellidse  of  Kansas,  the  writer 
accepts  the  conclusions  of  Funkhouser  and  others  in  making  the 
membracids  the  lowest  of  these  three  families  and  the  cercopids  the 
highest.    This  relationship  would  therefore  be  expressed  as  follows: 


FULGORIDAE 


CERCOPIDAE 


CICADELLIDAE 


MEMBRACIDAE 


CICADIOAE 


In  the  above  figure  it  will  be  noticed  that  the  Cercopidae  are  rep- 
resented as  branching  off  earlier  from  the  median  stem  than  the  other 
two  families.  This  seems  evident  when  their  life  history  is  consid- 
ered, for  they  have  been  isolated  long  enough  to  enable  them  to  per- 
fect a  method  of  protection  against  parasitism.  Certainly  the  devel- 
opment of  the  production  of  the  spittle  mass  which  incloses  the 
nymph  could  not  have  occurred  quickly,  but  must  have  taken  a 
relatively  long  period  of  time  to  attain  its  present  perfection. 

LIFE  HISTORY. 

Most  of  the  members  of  this  family  occur  on  small  trees  or  shrubs, 
particularly  those  growing  near  the  edges  of  woods  or  out  in  the 
open.    In  the  main  they  seem  to  avoid  the  shade  and  prefer  those 


42  The  University  Science  Bulletin. 

situations  where  they  receive  the  most  sunshine.  Most  of  our 
species  occur  on  trees,  but  there  are  several  that  are  found  on 
shrubs^  and  some  occur  on  weeds,  grasses  and  clovers. 

The  majority  of  our  species  of  tree  hoppers  are  found  to  over- 
winter in  the  egg  stage.  This  stage,  therefore,  occupies  by  far 
the  longest  period  in  the  life  history  of  the  insect.  The  eggs  are 
deposited  for  the  most  part  in  the  twigs.  Here  they  may  be  placed 
in  simple  incisions  in  a  row,  or  the  well-known  method  of  oviposi- 
tion  of  Ceresa  biibalus  may  be  used  where  two  curved  iuQisions, 
facing  each  other,  are  made.  Some  species  also  oviposit  in  the 
buds,  while  our  common  Entylia  concisa  lays  its  eggs  in  the  veins 
of  the  leaf  of  Ambrosia  trifida.  Funkhouser  gives  the  axils  of  the 
leaves  as  the  place  where  Telamona  ampelopsidis  oviposits,  while 
Thelia  bimaculata  lays  most  of  its  eggs  in  the  roots  or  on  the  stem 
below  the  surface  of  the  soil. 

While  most  of  our  treehoppers  hibernate  in  the  egg  stage,  Funk- 
houser states  that  Entylia  bactriana  and  Publilia  concava  over- 
winter as  adults.  The  writer  has  taken  Entylia  concisa  late  in  the 
fall  in  the  rubbish  and  grass  around  its  host  plant.  In  fact,  on 
September  8  he  has  found  the  eggs  of  this  species  just  hatching, 
while  on  September  11  he  has  taken  all  the  nymphal  instars  and 
newly  matured  adults  on  a  single  leaf. 

The  number  of  generations  in  a  season  is  usually  one,  by  far  the 
larger  number  of  our  species  having  a  single  annual  brood.  Some 
species,  however,  have  two  broods  and  some  probably  three,  while 
Vanduzea  arquata  is  said  by  Funkhouser  to  have  as  many  as  four 
generations.  The  same  is  seemingly  true  of  our  Vanduzea  trigut- 
tata. 

The  usual  life  history  of  a  tree  hopper  has  been  summarized  by 
Doctor  Funkhouser  as  follows: 

Eggs:   Laid  in  fall,  hatch  in  early  spring. 

Nymphs:  Emerge  about  the  middle  of  May  and  require  about  six  weeks 
to  reach  maturity. 

Adults:   Are  common  about  July  1  and  persist  throughout  summer  and  fall. 

Mating:   Takes  place  the  first  week  after  emergence. 

Oviposition :   Occui"s  within  a  week  after  mating. 

Broods:  Usually  one  but  sometimes  more,  dependent  on  weather  condi- 
tions. 


Lawson:    Membkacid.f.  of  Kansas.  43 

For  a  single  individual  the  life  cycle  would  be  somewhat  as  follows: 

Egg  stage:    From  SenleiiihiT  to  middle  of  May 8V^   months 

First   instar    1  week 

Second    instar    1  week 

Third    instar    1  week 

Fourth    instar    1  week 

Fifth  instar    2  weeks 

Nymph : 

Total,   fniiii  middle  of  May  to  July 1%   months 

Adult :    From  July  to  October  (inclusive)    4       months 

Entire  life    14       months 

The  above  dates  of  course  apply  to  New  York  and  would  differ 
somewhat  from  what  occurs  in  Kansas. 

One  of  the  easiest  ways  to  find  certain  species  of  tree  hoppers  is 
to  notice  the  more  readily  seen  ants  which  attend  so  many  of  our 
common  forms.  In  this  way  Vanduzea  triguttata  and  Entylia  con- 
cisa  are  very  easily  located.  It  is  well  known  that  the  ants  obtain 
honey  dew  from  these  insects,  although  there  are  many  species  of 
tree  hoppers  which  do  not  seem  to  furnish  this  substance  in  suffi- 
cient ciuantity  to  attract  the  attention  of  these  visitors. 

It  is  well  known  that  these  insects  are  frequently  parasitized. 
Hymeqopterous  parasites  of  the  family  Dryinidae  have  been  known 
to  occur  in  several  genera  of  tree  hoppers.  Doctor  Kornhauser 
worked  out  in  great  detail  the  life  history  of  such  a  parasite, 
Aphelopiis  thelice,  which  parasitizes  Thelia  himaculata.  In  addi- 
tion, tree  hoppers  are  often  the  prey  of  predaceous  insects  and 
spiders.  Of  such  predaceous  enemies  there  are  a  number  of  kinds, 
but  it  is  quite  doubtful  if  these  enemies  are  a  very  vital  factor  in 
the  natural  control  of  the  members  of  this  family. 

ECONOMIC  IMPORTANCE. 

The  chief  damage  done  by  tree  hoppers  is  by  their  oviposition. 
The  records  of  injury  to  young  orchards,  especially  to  apple,  are 
many.  The  type  of  oviposition  that  may  really  be  serious  is  that 
represented  by  the  egg  punctures  of  Ceresa  bubalus,  and  this  species 
is  seemingly  the  most  often  accused  of  all  the  species  concerned. 
The  writer  has  seen  young  orchards  in  which  the  outer  twigs  were 
all  roughened  up  by  the  oviposition  scars,  and  such  cases  are  numer- 
ous in  the  literature.  The  exact  damage  done,  however,  is  prob- 
lematical, many  inclining  to  the  opinion  that  as  a  rule  the  damage 
done  is  not  serious.  Others  hold  that  in  after  years  such  scars  will 
weaken  the  branches,  causing  them  to  be  more  easily  broken  off  by 
the  wind,  and  also  furnishing  good  points  of  attack  for  borers.    The 


44 


The  University  Science  Bulletin. 


wounds  made  also  doubtless  furnish  good  points  of  entrance  for  cer- 
tain fungous  diseases. 

Cases  are  on  record  also  where  the  buds  of  certain  trees  were 
seriously  injured  by  the  deposition  of  eggs  within  them.  Damage 
from  such  oviposition  always  results  when  eggs  are  deposited  deeply 
in  small  buds.  In  this  respect  the  species  concerned  most  fre- 
quently are  Ceresa  taurina,  Stictocephala  inermis  and  Enchenopa 
binotata.  The  species  most  injurious  to  stems  belong  to  the  genera 
Ceresa  and  Stictocephala. 

Inasmuch  as  the  nymphs  of  the  most  injurious  species  prefer  to 
feed  on  the  more  succulent  weeds  and  grasses  in  the  orchard,  the 
clean  culture  of  badly  infested  orchards  and  the  getting  rid  of 
near-by  weed  patches  in  the  early  summer,  will  usually  be  all  the 
control  measures  necessary.  Frequently  in  case  of  very  severe  in- 
festations the  careful  pruning  of  the  worst -infested  twigs  after  egg 
deposition  is  a  desirable  practice. 


LIST  OF  THE  SPECIES. 


Microceniru>i  caryce  (Fitch). 
Campylenchia  latipes  (Say). 
Enchenopa  binotata  (Say). 
Ceresa  diceros  (Say). 
Ceresa  albescens  Van  D. 
Ceresa  bubalus  (Fabr.) 
Ceresa  taurina  Fitch. 
Ceresa  palmeri  Van  D. 
Ceresa  borealis  Fairm. 
Ceresa  brevicornis  Fitch. 
Stictocephala  inermis  (Fabr.) 
Stictocephala  festina  (Say). 
Stictocephala  lutea  (Walk.). 
Acutalis  tartarea  (Say). 
Micrutalis  calva  (Say). 
Carynota  mera  (Say). 
Thelia  bimacnlata  (Fabr.). 
Thelia  uhleri  Stal. 
Glossonotus  acuminatus  (Fabr.) 
Heliria  cristata  (Fairm.). 
Heliria  s^alaris  (Fairm.). 
Telamona  pyramidata  (Uhl.). 
Telamona  viridia  Ball. 
Telamona  obsoleta  Ball. 
Telamona  decorata  Ball. 
Telamona  querci  Fitch. 
Telamona  ampelopsidis  (Harr.). 


Telamona  tristis  Fitch. 
Telamona  unicolor  Fitch. 
Telamona  extrema  Ball. 
Telamona  collina  (Walk.). 
Telamonanthe  rileyi  (Godg.). 
Telamonanthe  modesta  (Godg.). 
Archasia  galeata  (Fabr.). 
Archasia  belfragei  Stal. 
Smilia  camelus  (Fabr.). 
Cyrtolobus  celsus  Van  D. 
Cyrtolobus  juliginosus  (Emns.). 
Cyrtolobus  arcuatus  (Emns.). 
Cyrtolobus  jenestratus  (Fitch). 
Cyrtolobiis  tuberosus  (Fairm.). 
Cyrtolobus  I'au  (Say). 
Cyrtolobus  griseus  Van  D. 
Cyrtolobus  cinereus  (Emns.). 
Cyrtolobus  querci  (Fitch). 
Cyrtolobus  muticus  (Fabr.). 
Ophid^rma  salamandra  Fairm. 
Ophiderma  flaviguttida  Godg. 
Ophiderma  flava  Godg. 
Vanduzea  triguttata  (Burm.). 
Entylia  concisa  Walk. 
Pnblilia  concava  (Say). 
Publilia  modesta  Uhl. 
Publilia  reticulata  Uhl. 


Lawsox:    Membracid.T':  of  Kansas.  45 


SYSTEMATIC  TREATMENT  OF  KANSAS  SPECIES. 

The  three  subfamilies  represented  in  Kansas  may  be  separated 
by  the  following  key : 

KEY   TO    SUBFAMILIES. 

A.    Scutelhun  distinct,  not  concealed  by  the  pronotum.  Centrotinse. 
AA.   Sciitelhini  wanting  or  concealed  by  the  pronotum. 

B.    Tibiae,  especially  anterior  ones,  foliaceoiis.  Membracinae. 

BB.    Tibiae  not  foliaceoiis.  Smiliinae. 

Subfamily  CENTROTIN^   (Spinola). 

This  subfamily,  represented  in  the  United  States  by  about  half  a 
dozen  genera  and  a  few  more  species,  is  readily  separated  from  the 
other  subfamilies  by  the  presence  of  a  usually  visible  scutellum. 
The  genus  Microccntrus  occurs  in  Kansas. 

Genus  Microcentrus   (Stal). 

The  members  of  this  genus  that  occur  in  the  United  States  have 
a  pronotum  that  leaves  a  large  part  of  the  scutellum  well  exposed 
and  ends  in  a  slender  terminal  process  which  extends  beyond  the 
divided  apex  of  the  scutellum. 

Both  members  of  this  genus  known  to  occur  in  the  United  States 
have  been  taken  in  Kansas.  They  may  be  separated  by  the  follow- 
ing key : 

KEY   TO   SPECIES. 

A.    Prothorax  bearing  a  pair  of  laterally  flattened  horns.  perdita. 

AA.    Prothorax  without  laterally  flattened  horns.  caryce. 

Microcentrus  caryce  (Fitch). 

Uroxiphus  caryce  Fitch,  Hornop.  N.  Y.  St.  Cab.,  p.  52,  1851. 
Centrotus  caryce  Walker,  List  Homop.,  iv,  p.  1147,  1851. 
Microcentrus  caryce  Stal,  Of.  Vet.  Akad.  Forh.,  xvi,  p.  295,  1869. 
Piiaulocentrus  caryce  Fowler,  Biol.  Centr.  Am.,  Homop.,  ii,  p.  159,  1896. 

Funkhouser  gives  the  following  technical  description: 

Gray-brown  mottled  with  black;  entire  body  broad  and  flat;  pronotum 
roundly  swollen  above  line  of  abdomen  and  wings;  wings  broadJy  tectiform. 

Head  perpendicular,  twice  as  broad  as  long,  roughly  sculptured,  closely  jnmc- 
tate,  densely  pubescent,  deep  brown  at  base;  eyes  prominent,  extending  be- 
yond lateral  margins  of  pronotum,  dark  brown  margined  with  lighter;  ocelli 
small,  pearly,  farther  from  the  eyes  than  from  each  other,  with  deep  depression 
between  them;  clypeus  prominent,  broad,  lighter  in  color  than  vertex  above, 
extending  far  below  lateral  margin  of  head. 

Prothorax  subspherical,  with  high  median  carina,  coarsely  punctate,  pubes- 
cent ;  light  brown  marked  with  black  on  median  ridge  and  above  head ;  pos- 
terior margin  tnmcate  except  for  narrow  process,  which  projects  to  angles  of 


46  The  University  Science  Bulletin. 

tegmina  and  short  sharp  tooth  on  each  latero-posterior  angle.  Scutellum 
broadly  exposed,  wide  at  base,  truncate  at  tip,  which  does  not  reach  apex  of 
posterior  process. 

Tegmina  translucent,  pubescent,  inner  margins  straight  and  meeting  at 
median  dorsal  line;  veins  prominent  and  nodulate;  apices  of  tegmina  extend- 
ing beyond  tip  of  abdomen.  Legs  and  undersurface  of  body  light  brown 
mottled  with  white.    Undersurface  of  abdomen  often  tomentose. 

Length,  9-10  mm.;  with,  3  mm. 

Distribution.  Van  Duzee  records  this  species  from  Ontario,  New 
York,  New  Jersey,  Pennsylvania,  North  Carolina  and  Ohio.  In  the 
Crevecoeur  collection  is  a  specimen  taken  in  Pottawatomie  county, 
Kansas,  thus  establishing  its  occurrence  in  the  state. 

Hosts.  As  indicated  by  the  specific  name,  this  species  occurs  on 
hickory.  Funkhouser  reports  it  on  oak  also.  Matausch  gives  swamp 
oak  and  walnut,  and  Goding  mentions  the  latter. 

Microcentrus  perdita  (Amyot  &  Serville). 

Ledra  perdita  Amyot  &  Serville,  Heinip.,  p.  577,  pi.  11,  fig.  5,  1843. 
Centruchus  leibecki  Goding,  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  471,  1894. 
Microcentrus  perdita  Baker,  Can.  Ent.,  xxix,  p.  38,  1897. 
Centruclioides  perdita  Van  Duzee,  Bui.  Buf.  Soc.  Nat.  Sci.,  ix,  p.   117,  1908. 

Goding  describes  this  species  as  follows: 

Yellow-ferruginous,  silky  white  between  lateral  horns;  behind  horns  a  spot 
on  costal  margin,  near  base  of  tegmina.  Head  black,  broad,  eyes  verj'  prom- 
inent; base  convex,  griseous,  lower  part  of  face  strongly  declivous;  four  rough- 
ened caringe  pass  along  face  from  base  do\vnward,  the  ocelli  being  in  the  two 
internal  ones,  the  outer  ones  being  contiguous  to  the  ej'es;  part  of  face  below 
eyes  triangular,  apex  yellow.  Ocelli  equidistant  from  each  other  and  the  eyes. 
Prothorax  convex,  lateral  angles  slightly  produced,  a  prominent  median  carina 
extending  from  base  to  apex  nearly  black.  Above  the  lateral  angles,  on  each 
side,  is  a  long  horn  or  protuberance,  flattened  laterally,  slightly  curving  upward, 
outward  and  forward,  the  apex  truncated;  width  of  base  and  apex  equal.  Apex 
of  scutellum  bidentate,  the  teeth  ivory-white;  posterior  margin  of  prothorax 
with  a  very  slender  tooth  or  style,  extending  backward  on  each  side  of  pos- 
terior process  of  prothorax,  a  little  distant  from  it.  The  entire  surface  of  the 
prothorax  densely  and  regularly  punctured.  Apex  of  tegmina  far  surpassing 
tip  of  abdomen;  a  black  spot  on  the  internal  margin  a  short  distance  from  the 
apex;  another  black  spot  on  the  costa,  about  one-third  the  distance  from  the 
base.  Tarsi  black;  legs  mottled  with  ferruginous  and  grayish-yellow;  tibiae 
triquetrous.    Tegmina  lightly  ferruginous  and  opaque. 

Length  to  apex  of  tegmina,  8.5:  mm.;  width  at  lateral  angles,  3.2  mm. 

Distribution.  Reported  from  Massachusetts,  New  Jersey,  Penn- 
sylvania, Florida,  Colorado  and  New  Mexico.  Specimens  have  also 
been  taken  in  Riley  county,  Kansas. 

Hosts.    Van  Duzee  reports  this  specief>  from  scrub  oak. 


i 


Lawson:    Membracid.e  of  Kansas.  47 

Si-BFAMTLY  I\I KMBRACIX.E  (Stal). 

The  members  of  this  subfamily  differ  from  all  other  mcmbracids 
of  our  fauna  by  having  the  prothoracic  and  mesothoracic  tibiae 
foliaceous.  As  in  the  following  subfamily,  the  pronotum  conceals 
the  scutellum. 

Two  genera  occur  in  Kansas  ^^■ilich  may  be  separated  by  the  fol- 
lowing key: 

KEY   TO   GEiN'ERA. 

A.    Lateral  ridges  of  anterior  horn  n(\irer  dorsal  carina;  ventral  carina  of  horn 
not  distinctly  foliaceous.  Campylenchia. 

AA.   Lateral    ridges    of   anterior    horn    midway    between    dorsal    and    ventral 
carinse;  latter  distinctly  foliaceous.  Enchenopa. 

Genus  Campylenchia  (Stall. 

This  genus,  the  chief  characteristics  of  which  appear  in  tlie  above 
key,  is  represented  by  a  single  though  widely  spread  species  in  the 
United  States. 

Campylenchia  latipes  (Say). 

(PI.  VI,  figs,  9,  10.) 

Membracis  latipes  Say,  Long's  2nd  Exped.,  ii,  p.  302,  1824;    Conipl.  Writ.,  i,  p.  202. 

Enchophylluvi  latipes  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  47,  1851. 

Enchenopa  latipes  Walker,  List  Homop.,  ii,  p.  482,  1851. 

Enchenopa  antonina  Walker,  List  Homop.,  ii,  p.  488,  1851. 

Enchenopa  venosa  Walker,  List  Homop.,  ii,  p.  488,  1851. 

Enchenopa  frigida  Walker,  List  Homop.,  ii,  p.  490,  1851. 

Enchenopa  himacula  Walker,  List  Homop.,  ii,  p.   491,   1851. 

Campylenchia  curvata  Stal,  Heniip.  Fabr.,  ii,  p.   43,  1869. 

Enclienopa  curvata  Uhler,  Bui.  U.  S.  Geol.  Geog.  Surv.,  i,  p.  343,  1876;    iii,  p.   457,   1877. 

Aconophora  curvata  Butler,  Cist.  Ent.,  ii,  p.  349,   1877. 

Echenopa  rectidorsian  Buckton,   Monog.    Membr.,   p.    49,   1903. 

Campylenchia  latipes  Van  Duzee,  Can.  Ent.,  xliv,  p.  320,   1912. 

Funkhouser  gives  the  following  technical  description: 

Uniform  cinnamon  brown,  densely  punctate,  sparingly  pubescent;  single 
porrect  pronotal  horn  projecting  forward  over  head;  head  and  first  two  pairs 
of  legs  broadly  foliaceous,  hind  legs  spined;  tegmina  opaque,  punctate  at  basal 
and  costal  margins. 

Head  quadrate,  somewhat  declined,  shining  brown  somewhat  mottled  with 
darker,  lightly  punctate,  densely  pubsecent ;  eyes  prominent;  ocelli  small, 
pearly,  equidistant  from  each  other  and  from  the  ej'es  and  situated  on  a  line 
passing  through  centers  of  eyes;  clypeus  very  broad,  shining,  scarcely  punctate, 
broadly  truncate  at  apex,  tip  strongly  pubescent. 

Prothorax  produced  anteriorly  into  a  long,  flattened  horn,  ridged  in  center 
and  foliaceous  above  and  below,  varying  greatly  in  length  and  degree  of  curve; 
posterior  process  strong,  tectiform,  reaching  internal  angles  of  tegmina; 
median  dorsal  carina  strong  and  percuiTent ;  entire  pronotum  concolorous, 
lightly  punctate,  sparingly  pubescent  with  golden  hairs;  median  lateral  ridge 
reaching  lateral  margin. 

Tegmina   yellow-opaque;   basal  and   costal   areas   punctate   and  pubescent; 


48  The  University  Science  Bulletin. 

veins  distinct,  broad,  and  slightly  pubescent ;  fi\e  apical  and  two  discoidal 
cells;  hind  wings  iridescent.  Two  anterior  pairs  of  legs  broadly  spatulate  and 
lightly  pubescent  at  margins;  posterior  tibiae  armed  with  black-tipped  spines; 
tarsi  much  produced  and  lighter  in  color.  Undersurface  of  body  chocolate 
brown. 

Length:  from  head  to  apices  of  elytra,  5  mm.;  from  tip  of  promotal  horn 
to  apices  of  elytra,  8  mm.    Width  between  humeral  angles,  2  mm. 

Internal  male  genitalia.  Styles  with  long  flattened  anterior  proc- 
ess, widest  apically,  with  a  distinct  prominence  at  attachment  to 
connective,  caudad  of  which  they  are  peculiarly  wrinkled  and  show 
a  small  but  distinct  mesal  lobe,  then  widest  just  caudad  of  wrinkled 
portion,  after  which  they  curve  laterad,  ending  in  a  doubly  pointed 
apex,  of  which  the  terminal  tooth  is  larger  and  more  slender  than 
the  anterior  one,  the  apical  third  of  the  styles  bearing  a  few  short 
hairs;  connective  short  and  wide,  basal  half  with  a  heavily  chitinized 
and  flattened  V-shaped  portion;  oedagus,  viewed  laterally,  stout  and 
broadly  V-shaped,  widest  at  the  distinctly  notched  base,  the  upper 
basal  process  also  slightly  notched,  narrowest  at  the  bend  and  then 
widening  till  near  the  apex,  the  caudo-dorsal  portion  of  which  is  only 
slightly  chitinized,  the  functional  orifice  being  at  about  the  middle 
of  this  slightly  chitinized  area,  the  upper  portion  of  the  apex  bearing 
a  few  distinct  file-like  teeth. 

Distribution.  Van  Duzee  gives  the  following  distribution  for  this 
species:  Ontario,  Maine,  Vermont,  New  York,  New  Jersey,  Penn- 
sylvania, North  Carolina,  Illinois,  Iowa,  Kansas,  Missouri,  Colo- 
rado, Wyoming,  Montana,  Texas,  New  Mexico,  Arizona,  and  Cali- 
fornia. Osborn  reports  it  from  Florida.  It  is  thus  seen  to  extend 
throughout  the  United  States. 

In  Kansas  specimens  have  been  taken  in  the  following  counties: 
Douglas,  Ottawa,  Pratt,  Hodgeman,  Clark,  Pottawatomie,  Labette, 
Bourbon,  Wabaunsee,  Russell.  Riley,  Ellis,  Ellsworth  and  Wyan- 
dotte. 

Hosts.  Funkhouser  gives  the  following  hosts  for  this  species: 
Aster,  daisy,  joe-pye  weed,  alfalfa,  prickly  lettuce,  wild  carrot. 
Miss  Branch  mentions  golden  rod  and  sensitive  rose.  Gillette  and 
Baker  report  it  from  Glycyrrhiza  lepidota,  Psoralea  tenuijiora  and 
Apocynimi  androsacniifolium.  The  writer  has  found  it  very  abun- 
dantly on  alfalfa. 

Genus  Enchenopa  Amyot  &  Serville. 
But  two  members  of  this  genus  are  known  to  occur  in  the  United 
States.    The  widely  distributed  E.  binotata  occurs  in  Kansas. 


i 


Lawson:    Membracid.e  of  Kansas.  49 

Knchcnopa  binotnfa   (Say). 
(I'l.  \i,  ti«s.  n,  12.) 

Moiibracis  biiiotata  Say,   Long's  2nd  E.\ped.,   ii,  p.   3(ll,   1824;    Conipl.   Wril.,   1,  p.   201. 

Enchophyllum  binotatum  Fitch,  Homop.   N.   Y.   St.   Cab.,   p.    47,    18.51. 

Enchenopa  biiiotata  Walker,  I.,ist  Homop.,  ii,  p.  481,   1851. 

Encheiiopa  brevis  Walker,  List  Homop.,  ii,  p.  492,  1851. 

Thelia  biiiotata  Emmons.  Nat.  Hist.  N.  Y.  Ins.,  p.   15f>,   1854. 

Enchenopa  bifusifera  Walker,  List  Homop.,  f<uppl.,  p.  125,  1858. 

Enchenopa  bivittata  Rathvon  in  Mombert's  Hist.  Lane.  Co.,  Pa.,  p.   551,   1869. 

Enchenopa  porrecta  Buckton,  Monog.   Membr.,  p.   51,  pi.  (i,  fig.   5,   1902. 

Funkhouser  gives  the  following  technical  description: 

Much  resembling  the  preceding  species  in  size  and  in  general  appearance, 
but  differing  in  shape  of  the  head,  in  shape  of  sculpturing  of  the  pronotal 
horn,  and  in  bearing  two  yellow  spots  on  the  dorsal  line  of  the  pronotum. 

Head  longer  than  broad,  uniform  brown,  finely  but  densely  punctate,  spar- 
ingly pubescent;  eyes  prominent,  very  deep  brown;  ocelli  yellowish,  farther 
from  each  other  than  from  the  eyes;  clypeus  longer  than  broad,  rounded  at 
tip,  not  punctate. 

Prothorax  finel\-  i)unctate,  sparsely  pubescent;  two  distinct  ridges  on  each 
.side,  the  upper  extending  to  the  lateral  margin;  pronotal  horn  strongly  curved, 
broadly  foliaceous  above,  triquerate  at  tip;  median  dorsal  carina  high  and  per- 
ciu-rent;  two  dorsal  spots  of  lemon  yellow,  the  anterior  about  twice  as  long 
as  the  posterior;  posterior  process  gradually  acuminate,  extending  slightly 
beyond  internal  angles  of  tegmina. 

Tegmina  concolorous  brown,  opaque,  costal  margins  slightly  jjunctate,  and 
feebly  pubescent  at  base;  veins  distinct;  five  apical  and  one  discoidal  cell. 
First  two  pairs  of  legs  broadly  fohaceous;  hind  tibiae  spined;  tarsi  thin. 

Length  5  mm.;  width  2  mm. 

Internal  male  genitalia.  Styles  stout,  anterior  portion  longer  and 
broader  than  posterior,  with  a  rounded  prominence  to  connective, 
wider  just  caudad  of  this,  apices  strongly  hooked  and  truncate,  the 
apical  point  being  slightly  longer  than  the  anterior  one,  with  a  few 
hairs  on  the  terminal  third;  connective  heart-shaped,  broad,  a  basal 
V-shaped  band  heavily  chitinized;  cedagus,  viewed  laterally, 
strongly  curved,  base  very  wide  and  extending  dorsad  in  a  large 
rounded  prominence  which  is  separated  by  a  broad  notch  from  the 
pointed  attachment  to  the  connective,  the  apex  pointed  and  with 
functional  orifice  below  the  tip,  the  dorsal  surface  of  which  bears 
a  number  of  filelike  teeth. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  Mas- 
sachusetts, New  Hampshire,  New  York,  New  Jersey,  Pennsylvania, 
Maryland,  District  of  Columbia,  North  Carolina,  Florida,  Illinois, 
Iowa,  Michigan,  Kansas,  Missouri  and  Texas.  In  Kansas  speci- 
mens are  reported  from  Douglas,  Riley,  Sedgwick,  Bourbon  and 
Pottawatomie  counties. 


50  The  University  Science  Bulletin. 

Fowler  records  it  from  Mexico,  Guatemala  and  Panama.  It  is 
thus  seen  to  occur  throughout  tlie  eastern  United  States  and  down 
through  Central  America. 

Hosts.  Funkhouser  gives  the  following  hosts:  Locust,  wild  grape, 
bittersweet,  hickory,  sycamore,  butternut,  dogwood,  daisy,  joe-pye 
weed.  Miss  Branch  adds  golden  rod  and  pin  oak.  Matausch  gives 
Viburnum.  Goding  records  it  from  butternut,  birch,  apple,  walnut, 
grape,  hop  tree,  locust,  redbud,  cherry.  Viburnum,  Ceanothus  and 
white  birch.  The  writer  has  taken  it  chiefly  on  locust,  Ceanothus 
and  walnut. 

Subfamily  SMILIINtE  (Stal). 

This  subfamily  includes  all  but  the  three  preceding  members  of 
our  fauna.  In  all  of  them  the  pronotum  conceals  the  scutellum  and 
the  tibiae  are  not  foliaceous.  The  seventeen  genera  occurring  in 
Kansas  may  be  separated  by  the  following  key: 

KEY   TO   GENERA. 

A.   Tegmina  entirely  free,  not  covered  by  pronotum. 
B.   Veins  of  corium  closely  united  at  base. 

C.    Suprahumeral  horns  present.  Ceresa. 

CC.   Suprahumeral  horns  absent.  Stictocephala. 

BB.   Veins  of  corium  widely  separated  at  base. 

C.   Tegmina  with  five  apical  cells;  veins  distinct.  Acutalis. 

CC.   Tegmina  with  four  apical  cells;  veins  indistinct. 

Micrutalis. 
AA.   Tegmina  partly  or  entirely  covered  by  pronotum. 

B.   Terminal  cell  of  hind  wing  sessile,  its  base  truncate. 

C.    Pronotum  without  horn  or  crest. 

D.   Dorsum  low  and  rounded.  Carynota. 

DD.    Dorsum  high,  compressed,  and  foliaceous.      Archasia. 

CC.    Pronotum  with  horn  or  crest. 

D.    Horn  anterior  and  porrect.  Thelia. 

DD.    Horn  a  flat  dorsal  crest. 

E.    Crest  arising  from  between  humeral  angles. 

Glossonotus. 

EE.    Crest  arising  from  behind  humeral  angles. 

F.   Crest  strongly  step-shaped.  Heliria. 

FF.    Crest  sometimes  slightly,  but  not  usually  step- 
shaped. 

G.   Basal  costal  cell  of  tegmina  not  entirely 
punctate.  Telamona. 

GG.   Basal    costal    cell    of    tegmina    entirely 
punctate.  Telamoncmthe. 

BB.   Terminal  cell  of  hind  wing  triangular  and  petiolate. 

C.    Base  of  corium  with  three  veins. 

D.    Corium  without  cross-veins  at  base.  Smilia. 

DD.    Corium  with  cross-veins  at  base. 

E.    Dorsum  strongly  compressed.  Cyrtolobus. 

EE.   Dorsum  rounded.  Ophiderma. 


La\vso.\  :    Membracid^  of  Kansas.  51 

CC.   Base  of  corium  with  two  veins. 

D.    Apical  cell  of  te^;inina  transverse.  Vanduzea. 

DD.    Apical  cell  of  tegmina  triangular. 

E.    Dorsum  strongly  elevated,  with  deep  median  notch. 

Entylia. 

EE.    Dorsum   slightly   elevated,  with   weak   median  de- 
pression. Publilia. 

Genus  Ceresa  Amyot  &  Serville. 

To  this  genus  belong  our  commonest  species,  most  of  which  are 
green  and  all  of  which  are  at  once  recognized  by  their  prominent 
suprahumeral  horns. 

The  seven  species  known  to  occur  in  Kansas  may  be  separated  by 
the  following  key: 

KEY    TO    SPECIES. 

A.   Species  brown. 

B.    Species  larger  and   very   hairy;    suprahumerals   not   recurved;    one 
white  band  on  j)osterior  process.  diceros. 

BB.    Species  smaller,  with  few  hairs;  suprahumerals  recurved;  two  white 
bands  on  posterior  process.  albescens. 

AA.   Species  green. 

B.   Dorsal  crest  marked  with  brown  or  reddish. 

C.   Suprahumerals  short  and  slightly  recur\ed.  palmeri. 

CC.   Suprahumerals  longer  and  strongly  recurved.  con-stans. 

BB.    Dorsal  crest  concolorous. 

C.    Species  small,  7-8  mm.  long. 

D.   Very  hairy  species.  borecdis. 

DD.    Sparsely  haired  species.  brevicornis. 

CC.   Species  large,  8-10  mm.  long. 

D.    Suprahumerals  long,  sloping  upward  and  recurved;   cly- 
peus  much  produced  beyond  vertex.  taurina. 

DD.   Suprahumerals  stout,  nearly  straight,  clypeus  short. 

bubalus. 
Ceresa  diceros  (Say). 

(PI.  Ill,  figs.    ],   2.) 

Membracis  diceros  Say,  Long's  2nd  E.xped.,  ii,  p.   299,   1824;    Compl.  Writ.,  i,   p.   199. 

Smilia  diceros  Germar,   Silb.   Rev.   Ent.,  iii,   p.   237,   1835. 

Ceresa  postfasciata  Amyot  &  Serville,  Hem.,  p.  540,  pi.  10,  fig.  3,  1843. 

Ceresa  diceros  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.   2,  iv,  i).   285,  184fi. 

Ceresa  vitidalis  Buckton,  Monog.   Membr.,  p.   172,   pi.   36,  figs.   3-3b,   1903. 

Funkhouser  gives  the  following  technical  description: 

Dark  brown  with  trans\erse  bands  of  yellowish  white;  suprahumeral  horns 
stout  and  blunt;  posterior  process  decurved;  tegmina  smoky  hj'aline. 

Head  broader  than  long,  sculptured,  basal  part  strongly  and  smoothly 
curved,  front  surface  light  yellow  faintly  marked  with  brown,  faintly  longi- 
tudinally ridged,  very  slightly  or  not  at  all  punctate  or  pubescent ;  eyes 
prominent,  extending  beyond  adjoining  lateral  margin  or  pronotum;  ocelli 
shining,  transparent,  nearer  to  each  other  than  to  the  eyes;  sclerites  of  front 


52  The  University  Science  Billetin. 

projecting  over  clypeus  at  internal  angles  with  a  small  hook;  clypeus  strong, 
swollen,  roughly  three-Iobed,  the  central  lobe  the  largest,  tips  strongly  hirsute. 

Pronotuni  densely  and  coarsely  punctate;  anterior  surface  slightly  convex, 
light  yellow  with  numerous  brown  markings,  sparingly  pubescent  with  rather 
long  hairs;  suprahumeral  horns  projecting  outward  and  very  slightly  back- 
ward; lateral  surfaces  not  pubescent,  brown  with  two  transverse  light  bands, 
the  anterior  broad  and  irregular  in  about  center,  the  posterior  narrower  and 
regular  just  before  apex  of  j^osterior  process;  posterior  process  gradually  acute, 
extending  beyond  internal  angles  of  tegmina. 

Tegmina  hyaline,  tips  sriioky,  bases  opaque  and  lightly  punctate ;  five  apical 
and  three  discoidal  cells.  Undersurface  of  body  very  dark  brown.  Femora 
dark  brown  above;  tibise  and  tarsi  ferruginous. 

Length,  9  mm.;  width  between  humeral  horns,  5.5  mm. 

Internal  male  genitalia.  Styles  stout,  varying  much  in  the  length 
of  the  cephalic  part  which,  at  its  greatest  length,  is  shorter  than  the 
part  caudad  of  the  connective,  the  sharply  pointed  caudal  portion 
bearing  two  rows  of  long  hairs;  connective  longitudinally  keeled, 
when  open  nearly  twice  as  long  as  wide,  base  concave,  apex  nar- 
rowed but  obtuse;  oedagus,  viewed  laterally,  much  as  in  Ceresa 
bubalus,  but  with  dorsal  process  usually  more  pointed  apically  and 
the  ventral  process  distinctly  narrower  and  more  acute  apically. 

Distribution:  Van  Duzee  reports  this  species  from  Nova  Scotia, 
Ontario,  New  York,  New  Jersey,  Pennsylvania,  Maryland,  North 
Carohna,  Ohio,  lUinois,  Iowa,  Kansas,  Missouri,  Dakota,  Colorado, 
Texas,  New  Mexico  and  Montana.  It  is  therefore  one  of  our  most 
widely  distributed  species.  In  Kansas  specimens  have  been  taken  in 
Douglas,  Miami,  Neosho,  Bourbon,  Pottawatomie,  Riley,  Shawnee 
and  Saline  counties. 

Hosts:  The  usual  host  for  this  species  is  elderberry.  Funkhouser 
gives  the  following  additional  hosts:  Locust,  oak,  sycamore,  sweet 
clover,  blackberry  and  butternut. 

Ceresa  albescens  Van  Duzee. 

Ceresa  albescens  ^'an  Duzee,  Bui.  Buf.   Soc.  Nat.   Sri.,  ix.  p.   3.5,   1908. 
Ceresa  bubalus  var.  a  and  b  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.   50,  18.51. 

The  following  is  the  original  description: 

A  little  smaller  and  paler  than  diceros,  to  which  it  is  closely  allied.  Prono- 
tum  as  in  diceros,  but  with  the  suprahumeral  horns  more  acute  and  recurved 
and  tipped  with  black,  and  the  apex  longer  and  more  slender.  Face,  front 
and  superior  surface  of  the  pronotum  greenish-  or  yellowish-white  with  scarcely 
a  trace  of  the  maculations  found  in  dicero^i;  apex  of  the  head  less  produced, 
the  tylus  scarcely  longer  than  the  cheeks.  Sides  of  the  pronotum  paler,  fer- 
ruginous, becoming  somewhat  fuscous  posteriorly,  irrorate  with  paler  and 
marked  with  a  pale  marginal  line  and  sometimes  with  an  oblique  median 
vitta;  protracted  apex  whitish  with  a  black  tip  and  ferruginous  median  vitta; 
outer  surface  of  the  suprahumerals  dark  ferruginous,  differentiated  from  the 


Lawsox:    AlKMBRACiDiE  OF  Kansas.  53 

ferruginous  sides  by  the  pale  lateral  line  wliich  nearly  attains  the  apex  of  the 
horns.  Legs  and  beneath  t'(M-ruginous,  the  femora  darker.  Last  ventral  seg- 
ment of  the  female  a  \  tM y  little  oblique  and  roimded  to  the  median  notch, 
whirli  is  triangular.  acut(\  and  reaches  the  middle  of  the  segment.  Plates  of 
tlie  male  not  attaming  the  apex  of  the  anal  tube.  Elytra  almost  hyaline,  not 
smoky  brown  as  in  dircrus,  nervures  ferruginous.  Length,  8-9  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York,  and  Kansas.  Matauscli  took  specimens  in  New  Jersey.  Tlie 
Kansas  specimens  were  taken  at  Effingham,  Atchison  county. 

Hosts.    Matausch  gives  Viburnum  as  a  host. 

Ceresa  palmcri  Van  Duzee. 
(PI.  in,  fig^.  3,  4.) 

Ceresa  palmeri  \'aii  Duzee,  Can.   Ent.,  xl,   ii.    114,   1908. 

Funkhouser  gives  tlie  follo\ving  technical  description: 

Near  Ceresa  constarus,  but  differing  particularh-  in  shape  of  the  supra^ 
humeral  horns,  which  are  short,  terete,  and  but  little  recurved;  small,  reddish 
species,  with  jironotum  rather  high,  not  pubescent. 

Head  wider  than  long,  yellowish,  only  fain>tly  sculptured,  not  punctate; 
eyes  prominent,  reddish  with  white  borders,  extending  beyond  adjoining  lat- 
eral margins  of  pronotum ;  ocelli  not  prominent,  pearly  with  reddish  margins, 
nearer  to  each  other  than  to  the  eyes;  clypeus  continuing  lateral  margin  of 
face,  swollen  and  pubescent  at  tip. 

Pronotum  yellow-green,  very  strongly  marked  with  brown  and  reddish; 
dorsal  crest  curved,  strongly  marked  with  red;  lateral  semicircular  impression 
faint,  area  within  it  lighter  in  color  than  surrounding  pronotum;  posterior 
process  slightly  curved  downward,  about  reaching  tip  of  abdomen  but  not 
extending  halfway  to  extremities  of  tegmina. 

Tegmina  hyaline,  wrinkled,  bases  slightly  punctate.  L^ndersurface  of  body 
yellowish.  Legs  concolorous  3'ellow-green  in  life,  fading  to  pale  yellow  in 
cabinet  specimens. 

Length,  8  mm.;  width.  3.5  mm. 

Internal  male  genitalia.  Styles  large  and  stout,  widest  just 
caudad  of  attachment  to  connective,  then  broad  until  suddenly 
narrowed  to  the  acute  apex,  caudal  third  with  a  fringe  of  long 
hairs  extending  nearly  to  the  apex  on  the  outer  margin,  but  stop- 
ping considerably  cephalad  of  the  apex  mesally ;  connective  elongate 
heart-shaped;  cedagus,  viewed  laterally,  with  dorsal  process  of 
medium  width,  ventral  process  more  heavily  chitinized  and  quite 
stout  until  near  the  quite  acute  apex,  the  membrane  guarding  the 
functional  orifice  large  and  about  midway  between  the  base  and 
apex  of  the  process. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York  and  North  Carolina.  Specimens  have  been  taken  in  Kansas 
in  Linn,  Bourbon.  Hodgeman  and  Cowley  counties. 


54  The  University  Science  Bulletin. 

Hosts.  Funkhouser  gives  young  hickory  as  a  host,  while  Metausch 
took  numerous  nymphs  of  the  species  from  Liquidambar. 

Ceresa  constans  (Walker). 

Thelia  constans  Walker,  List  Homop.,  ii,  p.   563,   1851. 
Ceresa  constans  Stal,  Of.  Vet.  Akad.  Forh.,  xxvi,  p.  245,  1869. 
Ceresa  suhulata  Provancher,  Pet.  Faune  Ent.   Can.,  iii,  p.   3.38,   1890. 
Ceresa  illinoiensis  Goding,  Bui.  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  404,   1894. 

Funkhouser  gives  the  following  technical  description: 

Small  and  distinctly  reddL^ih;  dorsal  cre.st  low,  median  lateral  line  red;  meto- 
pidium  convex;  horns  gharji  and  much  recurved;  i)Osterior  process  nearly 
straight,  usually  tipped  with  red;  head  triangidar;  tegmina  and  wings  hyaline. 

Head  subtriangular,  weakly  sculptured,  faintly  longitudinally  furrowed,  very 
finely  and  lightly  punctate,  not  pubescent;  eyes  prominent,  dark  brown  with 
lighter  edges,  extending  beyond  adjoining  lateral  margins  of  pronotum;  ocelli 
glassy,  nearer  to  each  other  than  to  the  eyes;  clypeus  much  longer  than  wide, 
extending  for  more  than  half  its  length  beyond  lateral  margin  of  face,  tip 
hirsute. 

Pronotum  deeply  and  coarsely  punctate,  not  pubescent,  median  carina 
])rominent;  dorsal  crest  low,  rising  but  little  higher  than  tips  of  suprahumeral 
horns;  horns  slender,  sharp,  hiuch  recurved,  extending  upward  and  curving 
backward;  metopidium  convex,  regular;  lateral  semicircular  impression  deep, 
concolorous;  posterior  process  nearly  straight,  not  reaching  the  extremity  of 
the  abdomen  and  reaching  barely  one-third  the  distance  to  the  tips  of  the 
tegmina. 

Tegmina  hyaline.     Undersurface  of  body  and  legs  yellowish. 

Length,  8  mm.;  width,  4  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York,  North  Carolina  and  Illinois.  Specimens  have  also  been  taken 
in  Riley  county,  Kansas. 

Hosts.    Funkhouser  gives  locust  as  the  host. 

Ceresa  borealis  Fairmaire. 

(PI.  Ill,  figs.  7,  8.) 
Ceresa  borealis  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  284,  1846. 

Funkhouser  gives  the  following  technical  description: 

Resembling  C.  bubalus  in  general  outline,  but  much  smaller  and  very  hairy; 
metopidium  convex;  dorsum  curved,  posterior  process  only  slightly  decurved; 
head  impunctate;  notch  of  last  ventral  segment  of  female  broad  and  tri- 
angular. 

Head  broader  than  long,  yellowish,  roughly  sculptured,  faintly  longitudi- 
nally striate,  not  punctured  nor  pubescent;  eyes  prominent,  mottled  with 
green  and  brown,  extending  beyond  adjoining  lateral  margins  of  pronotum; 
ocelli  small,  reddish,  nuich  nearer  to  each  other  than  to  the  ej^es;  clypeus 
rounded,  somewhat  protruding,  extending  for  more  than  half  its  length  below 
lateral  margin  of  face,  tip  hirsute. 

Pronotum  green,  finely,  deeply  and  densely  punctate,  very  hairy;  meto- 
pidium convex;  median  carina  faint;  suprahumeral  horns  stout,  blunt,  nearly 


Lawson:    Membracid^  of  Kansas.  55 

straight,  projecting  almost  dircctlj'  outward;  dorsal  crest  regularly  arcuate; 
lateral  semicircular  inii)ression  nearly  obsolete;  posterior  process  curving 
slightly  downward,  not  extending  beyond  tip  of  abdomen  and  reaching  only 
for  a  short  distance  beyond  internal  angles  of  tegmina. 

Tegmina  entirely  hyaline,  somewhat  wrinkled,  bases  lightly  punctate.  Legs 
and  undersurfaces  of  bodj'  concolorous  greenish. 

Length,  S  mm.;  width,  4  mm. 

Internal  male  genitalia.  Style  very  large  when  compared  with 
the  otiier  members  of  the  genus,  anterior  process  short,  a  small  but 
distinct  knob  to  the  connective,  posterior  portion  extending  straight 
until  considerably  past  the  connective,  then  bending  rather  abruptl}^ 
laterad  and  ending  in  a  tip  that  on  its  mesal  margin  bears  a  number 
of  small  but  distinct  teeth;  connective  very  long  and  rather  slender, 
notched  at  base;  cedagus,  viewed  laterally,  small,  with  dorsal  proc- 
ess strongly  humped,  the  ventral  process  with  basal  knob,  then  ta- 
pering regularly  to  acute  tip,  membrane  around  functional  orifice 
at  middle  third. 

Distribution.  Van  Duzee  records  this  species  from  Ontario,  New 
York,  New  Jersey,  Pennsylvania,  North  Carolina,  Ohio,  Kansas, 
Colorado,  and  Utah.  In  Kansas  specimens  have  been  taken  in  the 
following  counties:  Cowley,  Sumner,  Bourbon,  Cherokee,  Chau- 
tauqua, Douglas,  Riley,  Shawnee,  Crawford,  Miami,  Linn,  Neosho, 
Rawlins,  and  Ottawa.    It  is  thus  seen  to  extend  well  over  the  state. 

Hosts.  Funkhouser  gives  the  following  host  list  for  this  species: 
Wild  grape,  locust,  elder,  willow,  oak,  hickory,  pignut,  raspberry, 
sycamore,  apple,  and  pear. 

Ceresa  brevicornis  Fitch. 

Ceresa  brevicornis  Fitch,  Trans.  N.  Y.  St.  Agr.  Soc,  xvi,  p.  451,  1856. 

The  following  is  the  original  description: 

This  is  so  similar  to  the  common  Buffalo  tree  hopper  that  it  will  scarcely 
be  distinguished  from  it  except  by  a  practiced  ej'e,  although  it  is  undoubtedly 
a  distinct  species.  It  differs  from  that  in  having  the  horns  much  more  short, 
and  the  sides'  of  the  thorax  when  viewed  in  front  are  not  gradually  curved  out- 
wards, but  are  straight  or  rectilinear,  with  the  horns  abruptly  projecting  from 
the  comer  at  the  upper  end  of  this  line.  The  acute  spine  at  the  tip  of  the 
thorax  is  also  more  long  and  slender.  The  thorax  between  the  horns  is  slightly 
convex.  The  dried  specimen  is  of  a  pale  dull  j-ellow  color  freckled  with  faint 
pale  green  dots  and  with  a  paler  straw-colored  stripe,  quite  distinct,  upon  the 
angular  sides  of  the  thorax  from  each  eye  upward  to  the  horn  and  from  thence 
to  the  summit  of  the  thorax. 

Length  of  the  female,  0.36. 

It  was  met  with  upon  hickory  bushes  in  New  Jersey. 


56  The  University  Science  Bulletin. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  Con- 
necticut, New  York,  New  Jersey,  Pennsylvania,  Ohio,  and  Illinois. 
A  single  specimen  taken  at  Kansas  City,  Mo.,  and  determined  by 
Dr.  E.  D.  Ball,  evidently  puts  this  species  within  our  range. 

Hosts.    Van  Duzee  gives  basswood  as  a  host. 

Ceresa  taurina  Fitch. 

(PI.   II,  figs.  3,  4.) 

Membracis  taurina  Harris,  List  Ins.  Mass.  in  Hitchcock,  Geol.  Mass.,  p.  579,  1833  (MS 
name). 

Enchenopa  taurina  Walker,  List  Homop.,  ii,  p.  495,  1851   (MS  name). 
Ceresa  taurina  Fitch,  Trans.  N.  Y.  St.  Agr.  Soc,  xvi,  p.   335,  1856. 

Funkhouser  gives  the  following  technical  description: 

Slightly  smaller  than  C.  bubalus,  but  resembling  it  in  color;  body  slender 
and  metopidiiim  conca\'e  transversely;  horns  sharp,  curving  upward  and  back- 
ward. 

Head  roughly  triangular,  wider  than  long,  roughly  sculptured,  not  punctate 
nor  pubescent,  basal  margin  strongly  curved;  eyes  prominent,  brown  and  in 
some  cases  barred  with  darker,  extending  beyond  the  adjoining  lateral  mar- 
gins of  the  pronotum;  ocelli  prominent,  pearly,  occasionally  margined  with 
reddish,  nearer  to  each  other  than  to  the  eyes;  clypeus  subrectangular,  swollen 
and  protruding,  extending  for  half  its  length  beyond  lateral  margin  of  face, 
faintly  trilobed,  apex  bristled. 

Pronotum  deeply  and  coarsely  pimctiu'ed,  bright  green  fading  to  yellow, 
sparingly  pubescent;  metopidium  strongly  concave,  with  curved,  transverse 
margin,  area  above  eyes  smooth;  suprahvmieral  horns  slender  and  sharp,  ex- 
tending upward  and  backward,  often  much  curved,  tips  generally  darker  than 
bases;  dorsal  crest  high  and  strongly  curved;  semicircular  lateral  impression 
deep  and  brownish;  posterior  process  slender,  strongly  decurved,  extending 
beyond  apex  of  abdomen  and  halfway  to  tips  of  tegmina. 

Tegmina  and  wings  entirely  hj'aline.  Underparts  of  body  and  legs  yellow- 
green. 

Length,  including  tegmina,  9  mm.;  width  between  tips  of  horns,  5.5  mm. 

Internal  male  genitalia.  Styles  with  anterior  part  weak,  strongly 
narrowed  near  middle  of  connective,  then  much  widened  into  the 
large  posterior  portion,  which  ends  in  an  oblique  and  serrate  tip, 
both  margins  of  the  caudal  half  of  the  posterior  portion  bearing 
long  hairs;  connective  long  and  rather  narrow,  folding  longitudi- 
nally, and  with  the  usual  basal  incision;  cedagus,  viewed  laterally, 
with  basal  process  long  and  slender,  with  a  distinct  knob  at  point  of 
attachment  to  the  connective,  the  ventral  process  rather  narrow, 
tapering  gradually  to  subacute  tip,  functional  orifice  covering  the 
middle  third  of  the  ventral  side. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  Mas- 
sachusetts, New  York,  New  Jersey,  Pennsylvania,  Virginia,  North 


Lawson:    Membracid^  of  Kansas.  57 

Carolina,  Ohio,  Michigan,  Iowa,  Kansas,  Colorado,  and  Arizona. 
Specimens  have  been  taken  in  Kansas  in  Sumner,  Hodgeman,  Doug- 
las, Riley,  Shawnee,  Ottawa,  Linn,  Cherokee,  Bourbon  and  Miami 
counties. 

Hosts.  Funkhouser  reports  this  species  from  the  following  hosts: 
Raspberry,  hickory,  potato,  blackberry,  dahlia,  hazelnut,  locust, 
witch  hazel,  blue  grass,  oak,  pear,  apple,  sweet  clover,  and  bitter- 
sweet. Miss  Branch  adds  horseradish  and  choke  cherry.  Matausch 
mentions  Solidago. 

Ceresa  bubalus  (Fabricius). 

(PI.  II,  figs.   1-2  and  5-8.) 

Membracif  bubalus  Fabricius,  Ent.  Syst.,  iv,  p.  14,   1794. 
Centrotus  bubalus  Fabricius,  Syst.  Rhyng.,  p.   20,   1803. 
Ceresa  bubalus  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  50,  1851. 

Funkhouser  gives  the  following  technical  description. 

Bright  green  fading  to  yellowish  in  cabinet  specimens;  horns  heavy  and 
stout,  pointing  directly  outward;  metopidium  broadly  convex;  dorsal  crest 
high  and  regularly  arched;  posterior  process  slender  and  recurved;  tegmina 
and  hind  wings  entirely  hyaline;  clypeus  heavy,  stout,  and  bristled. 

Head  one-third  broader  than  long,  longitudinal  striate  sculpturing;  basal 
part  broadly  curved,  front  surface  yellow,  not  punctate  nor  pubescent;  eyes 
prominent,  dark  brown,  extending  beyond  lateral  margin  of  pronotum  adjoin- 
ing; ocelli  prominent,  protruding,  with  brilliant  orange  borders,  nearer  to  each 
other  than  to  the  eyes;  clypeus  strong,  heavy,  continuing  lateral  outline  of 
face,  apex  bristled. 

Pronotum  densely  and  coarsely  punctate;  metopidium  strongly  convex, 
smooth  impunctate  areas  above  the  eyes,  sparingly  pubescent  with  short  scat- 
tered hairs;  suprahumeral  horns  stout,  blunt,  projecting  almost  directly  out- 
ward, not  at  all  upward,  tips  often  brownish,  whitish  line  extending  backward 
from  tip  to  lateral  margin;  lateral  surface  marked  with  light-colored  semicir- 
cular impression;  posterior  process  slender,  depressed,  extending  half  way  to 
apices  of  tegmina  and  slightly  beyond  tip  of  abdomen,  apex  brownish. 

Tegmina  hyahne,  bases  lightly  punctate.  Undersurface  of  body  yellowish. 
Legs  greenish. 

Length  to  apices  of  tegmina,  10  mm.;  width  between  horns,  6  mm. 

Internal  male  genitalia.  Styles  seemingly  varying  considerably 
in  size,  usually  large,  the  anterior  portion  about  as  long  as  the 
posterior,  which  ends  in  a  sharp  point  which  is  usually  perfectly 
smooth,  but  sometimes  bears  a  few  indistinct  teeth,  the  margins 
with  two  rows  of  hairs  of  which  the  outer  one  is  longer  and  consists 
of  longer  hairs;  connective  about  oval  in  outline,  the  base  incised 
and  with  a  distinct  tendency  to  fold  along  a  longitudinal  keel; 
oedagus,  viewed  lateral!}',  wdth  the  dorsal  process  varying  from 
rather  slender  and  straight  to  quite  stout  and  hunch-backed,  the 


58  The  University  Science  Bulletin. 

ventral  process  of  medium  thickness,  the  apex  acute  and  with  the 
usual  membrane  around  the  functional  orifice  near  the  middle  third. 

Distribution.  This  species  occurs  throughout  the  United  States 
as  shown  by  the  following  records  of  its  distribution  given  by  Van 
Duzee:  Nova  Scotia,  Ontario,  New  Hampshire,  Massachusetts,  New 
York,  New  Jersey,  Pennsylvania,  Maryland,  Delaware,  North  Caro- 
lina, Tennessee,  Ohio,  Indiana,  Illinois,  Iowa,  Minnesota,  Kansas, 
Missouri,  Dakota,  Montana,  Wyoming,  Colorado,  Utah,  New  Mex- 
ico, Texas,  and  California. 

As  far  as  our  collecting  in  this  state  shows,  this  species  is  by  far 
the  most  widely  distributed,  specimens  having  been  taken  in  the 
following  counties:  Douglas,  Wilson,  Riley,  Comanche,  Russell, 
Butler,  Pawnee,  Finney,  Graham,  Rawlins,  Cherokee,  Wallace, 
Harper,  Shawnee,  Wyandotte,  Allen,  Decatur,  Harvey,  Saline,  Neo- 
sho, Sumner,  Reno,  Hodgeman,  Osborne,  Labette,  Anderson,  Potta- 
watomie, Wabaunsee  and  Ottawa. 

Hosts.  The  list  of  plants  on  which  this  species  feeds  is  very 
large,  including  many  weeds  and  trees.  Funkhouser  records  it 
from  the  following  hosts:  Sycamore,  aster,  poplar,  potato,  butter- 
nut, hazelnut,  pear,  sumac,  oak,  locust,  elm,  willow,  elder,  sweet 
clover,  hickory,  pignut,  apple.  Miss  Branch  reports  it  from  osage 
orange,  horseradish,  gama  grass,  sunflower  and  alfalfa.  Coding 
records  it  from  apple,  potato,  tomato,  pear,  peach,  plum,  grape, 
apricot,  almond,  willow,  locust  and  Japanese  lily.  Gillette  and 
Baker  report  it  from  willow,  apple,  soft  maple,  Solidago  spectabilis, 
Aster  canescens,  Apocynum  androscemijolium,  alfalfa,  and  Glycr- 
rhiza  lepidota.  The  writer  has  taken  it  most  frequently  on  apple, 
locust,  hickory,  sweet  clover  and  alfalfa  in  Kansas. 

.  Genus  Stictocephala  Stal. 

The  members  of  this  genus  ai^e  greenish  species,  which  differ  from 
those  of  the  preceding  genus  by  lacking  the  suprahumeral  horns. 

The  three  species  occuring  in  Kansas  may  be  separated  by  the 
following  key: 

KEY  TO   SPECIES. 

A.    Carinate  sides  of  the  metopidium  meeting  before  the  middle  of  the  body. 

iiiermis. 
AA.    Carinate  sides  of  the  metopidium  meeting  at  or  behind  the  middle  of 
the  body. 
B.    Carinate  sides  of  the  metopidium   meeting  at  or  near  middle  of 
dorsum.  lutea. 

BB.    Carinate    sides    of    metopidium    meeting   much   behind    middle    of 
dorsum.  fcstina. 


Lawson:    Membracid.e  of  Kansas.  59 

Stictocephala  inermis  (Fabriciiis). 

(PI.  IV,  figs  1,  2,  and   5-8.) 

Menibracis  inermis  Fabricius,  Syst.  Ent.,  p.  G77,  1775. 

Cicada  inermis  Gmelin  in  Linnaeus,  Syst.  Nat.,  edn.  13,  i,  pt.  4,  p.  2093,  1788. 
Centrotus  inermis  Fabricius,  Syst.  Rhyng.,  p.  21,  1803. 

Membracis  goniphora  Say,  Jl.  Acad.  Nat.  Sci.  Phila.,  vi,  p.  243,  1830;  Compl.  Writ., 
ii,  p.  377. 

Smilia  inermis  Fitch,  Ilomop.   N.   Y.   St.   Cab.,   p.   48,   1851. 

Ceresa  gonophora  Walker,  List  Homop.,  iv,  p.  1141,  1851. 

Thelia  inermis  Walker,  List  Homop.,  iv,  p.  1142,  1851  (under  lutea). 

Stictocephala  inermis  Stal,  Of.  Vet.  Akad.  Forh.,  xxvi,  p.  246,  1869. 

Stictocephala  sanguino-apicalis  Coding,  Bui.  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  408,  1894. 

Funkhouser  gives  the  following  technical  (iescription : 

Fine  large  species,  brilliant  green,  slowly  fading  to  yellowish  in  dried 
material;  metopidium  perpendicular;  dorsal  crest  high  and  arcuate;  posterior 
process  slender  and  curving  downward;  tegniina  and  wings  entirely  hj'aline; 
upper  parts  of  femora  often  marked  with  black. 

Head  broad,  nearly  smooth,  very  finely  and  faintly  punctate,  longitudinally 
striate;  eyes  prominent,  subtriangular,  very  dark  bordered  with  white,  ex- 
tending beyond  adjoining  lateral  margins  of  pronotum;  ocelli  prominent, 
brownish,  nearer  to  each  other  than  to  the  eyes;  inferior  margins  of  vertex 
broadly  sinuate;  clypeiis  broad,  sparingly  pubescent,  median  lobe  of  apex 
extending  below  lateral  lobes. 

Pronotum  densely  and  coarsely  but  not  deeply  punctured ;  metopidium  con- 
vex, median  carina  distinct  but  irregular;  sides  of  metopidium  meeting  be- 
fore middle  of  body;  lateral  semicircular  impression  deep;  posterior  process 
long,  slender,  gradually  acuminate,  curving  downward,  extending  beyond  abdo- 
men and  reaching  about  halfway  from  internal  angles  to  apices  of  tegmina. 

Tegmina  entirely  hyaline,  slightly  wrinkled,  bases  greenish  and  lightly 
punctured.  Undersurface  of  body  j'ellowish;  segments  of  abdomen  in  some 
cases  bordered  with  black;  notch  of  last  ventral  segment  of  female  broadlj^ 
angular.    Femora  often  marked  with  black  above;  tarsi  ferruginous. 

Length  to  tips  of  tegmina,  9  mm.;  width  between  humeral  angles,  4  mm. 

Internal  male  genitalia.  Styles  large  anci  stout,  the  anterior  por- 
tion long  and  wicie,  the  posterior  part  curving  to  the  truncate  anci 
serrate  tips,  which  vary  from  nearly  transversely  truncate  to  quite 
obliquely  truncate,  the  apex  in  the  latter  case  being  quite  pro- 
nounced, the  posterior  fourth  with  two  rows  of  very  long  hairs,  the 
outer  row  being  longer;  connective  short,  widest  just  behind  the 
middle,  usually  pentagonal  in  shape;  oedagus,  viewed  laterally, 
much  as  in  Ceresa,  the  dorsal  process  varying  in  width  and  with  a 
small  to  a  very  pronounced  hump,  but  with  no  projection  to  the 
connective,  ventral  process  moderately  stout,  slightly  concave  on 
ventral  side  preapically,  the  functional  orifice  occupying  nearly  half 
its  length  medially. 

Distribution.    This  species  is  found  throughout  the  United  States 


60  The  University  Science  Bulletin. 

as  shown  by  the  following  distribution  giyen  by  Van  Duzee:  On- 
tario, New  York,  New  Jersey,  Pennsylvania,  North  Carolina,  Ohio, 
Illinois,  Iowa,  Dakota,  Kansas,  Missouri,  Colorado,  New  Mexico, 
Arizona,  California,  and  Montana. 

In  Kansas  it  has  been  taken  in  the  following  counties :  Chautau- 
qua, Douglas,  Hodgeman,  Cowley,  Ottawa,  Riley,  Dickinson,  Linn, 
Ellis,  Bourbon,  Montgomery,  and  Wabaunsee. 

Hosts.  Funkhouser  records  this  species  from  sweet  and  red 
clover,  timothy  and  apple.  Miss  Branch  mentions  gama  grass. 
Coding  reports  it  from  plum,  oats,  oak  and  alfalfa.  The  writer 
has  taken  it  very  commonly  on  apple. 

Stictocephala  lutea  (Walker). 

Thelia  lutea  Walker,  List  Homop.,  ii,  p.   559,   1851. 

Thelia  inermis  Walker,   List   Homop.,   iv,  p.   1142,   1851. 

Gargara  pectoralis  Emmons,  Nat.  Hist.  N.  Y.  Ins.,  p.  157,  p.  1,  13,  fig.  12,  1854. 

Stictocephala   lutea  Stal,  Of.   Vet.    Akad.    Forh.,   xxvi,  p.   247,   1869. 

Funkhouser  gives  the  following  technical  description: 

Small  species;  grass-green  above,  usually  marked  with  black  below;  meto- 
pidium  sloping,  dorsal  crest  not  high,  not  regularly  arcuate;  tegmina  smoky 
hyaline. 

Head  perpendicular,  subtriangiilar,  broader  than  long,  finely  punctate, 
sparingly  pubescent,  weakly  sculptured;  eyes  prominent,  brown  usually  banded 
with-  reddish,  extending  outward  as  far  as  lateral  angles;  ocelli  distinct,  yel- 
lowish margined  with  brown,  much  nearer  to  each  other  than  to  the  eyes; 
inferior  margins  of  vertex  weakly  sinuate,  their  ventral  raesal  angles  ending  in 
hooks;  clypeus  robust,  extending  only  slightly  beyond  inferior  margins  of 
vertex. 

Pronotum  closely  and  deeply  punctate;  metopidium  convex,  median  carina 
faint,  smooth  yellowish  area  on  each  side  near  base  of  head,  sides  of  meto- 
pidium meeting  at  or  a  little  before  middle  of  body;  dorsal  crest  not  high, 
sloping  gradually  from  junction  of  carinate  edges  of  metopidium  to  posterior 
process;  semicircular  lateral  impression  weak;  posterior  process  slender,  gradu- 
ally acute,  extending  as  far  as  tip  of  abdomen  and  to  a  point  on  tegmina  half- 
way between  internal  angles  and  apices. 

Tegmina  hyaline,  smoky  at  apices.  Under  parts  of  thorax  distinctly  black. 
Legs  generally  marked  with  black.  Notch  of  last  ventral  segment  of  female 
very  small  or  obsolete. 

Length,  6.5  mm.;  width,  2  mm. 

Internal  male  genitalia.  Styles  stout,  especially  posterior  portion, 
bent  in  near  middle  of  connective,  then  flaring  widely  till  just  before 
the  incurved  tips,  which  are  transversely  truncate  with  the  inner 
angle  prominent  and  distinctly  serrate  on  both  its  margins,  the 
apical  fourth  of  the  styles  bearing  on  each  margin  a  row  of  long 
hairs;  connective  large,  elongate,  widest  just  caudad  of  the  middle; 
cedagus,  viewed  laterally,  with  medium-sized  and  humped  dorsal 


J 


Lawson:    Membracid.e  of  Kansas.  61 

process,  the  ventral  process  with  a  distinct  tubercle  to  connective 
and  wide  till  the  end  of  the  functional  orifice,  then  suddenly  nar- 
rowed, after  which  it  is  nearly  parallel-margined  to  the  apex. 

Distribution.  Seemingly  most  abundant  in  the  eastern  United 
States  as  shown  by  the  following  records  given  by  Van  Duzee:  On- 
tario, Connecticut,  New  York,  New  Jersey,  Pennsylvania,  District 
of  Columbia,  North  Carolina,  Georgia,  Florida  and  Illinois.  There 
are  specimens  of  this  species  in  the  Snow  collection  from  Beaver 
Creek,  Montana,  thus  extending  its  range  westward. 

The  following  Kansas  counties  have  yielded  specimens:  Linn, 
Montgomery,  Neosho,  Riley  and  Douglas. 

Hosts.  Matausch  reports  this  species  from  Solidago;  Funkhouser 
from  oak  and  daisy ;  Coding  from  wheat. 

Stictocephala  festina  (Say). 

(PI.  HI,  figs.  5,  6.) 

Membracis  festina  Say,  Jl.  Acad.  Nat.  Sci.  Phila.,  vi,  p.  243,  1830;  Compl.  Writ.,  ii,  p. 
377. 

Stictocephala  festiva  Walker,  List  Honiop.,  iv,  p.  1141,  1852. 
Stictocephala  uniformis  Stal,  Hemip.  Fabr.,  ii,  p.  24,  1809. 
Stictocephala  festina  Stal,  Of.  Vet.  Akad.  Forh.,  xw-i,  p.  246,  1869. 

The  following  is  the  original  description: 

Thorax  with  a  subacute  Hne  each  side  before,  meeting  behind  the  middle. 

Inhabits  Florida. 

Body  yellowish-green;  thorax  unarmed,  carinate  behind;  at  tip  attenuated, 
subulate  and  complying  with  the  general  cunature ;  each  side  before  a  carinate 
line,  meeting  together  at  the  carina  behind  the  middle,  with  the  carina  tinged 
nith  rufous;  front  of  the  thorax  not  altogether  flat,  but  a  little  convex;  heme- 
lytra,  three  temiinal  cellules  unequal;  the  two  costal  ones  equal,  as  broad  as 
long;  the  inner  one  not  Qb\'iousl3-  larger  than  the  others  together,  somewhat 
longer  than  broad.  Length  to  tip  of  hemelytra  one-fifth  of  inch.  The  lateral 
prominent  lines  of  the  unarmed  thorax,  separate  this  species  from  all  those 
I  have  described  excepting  goniphera,  which  meet  before  the  middle  of  the 
length  of  the  back. 

Internal  male  genitalia.  Styles  quite  small,  anterior  portion 
smaller  than  posterior,  converging  posteriorly  to  the  wide  posterior 
portions,  which,  opposite  the  connective,  have  a  distinct  lateral  an- 
gle and  then  narrow  but  slightly  to  the  obtuse  apices,  each  bearing 
a  long  outer  row  and  a  short  mesal  row  of  stout  hairs;  connective 
quite  small  and  triangular;  cedagus,  viewed  laterally,  quite  charac- 
teristic, the  dorsal  process  ending  in  a  large  rounded  lobe  which 
bears  a  small,  fingerlike  terminal  process,  the  ventral  process  ending 
in  a  swollen  and  obliquely  truncate  apex  which  bears  the  functional 
orifice. 


62  The  University  Science  Bulletin. 

Both  the  styles,  connective  and  oedagus  of  this  species  are  so  en- 
tirely different  from  the  corresponding  parts  in  the  two  preceding 
species  that  it  does  not  seem  possible  that  they  could  be  members 
of  the  same  genus. 

Distribution.  This  species  occurs  abundantly  in  the  Southern 
states  and  in  more  limited  numbers  as  far  north  as  Connecticut,  and 
Canada  in  the  East  and  Montana  in  the  West.  It  has  been  reported 
from  Florida,  Virginia,  Pennsylvania,  Georgia,  Missouri,  Texas, 
Iowa,  Montana,  Colorado,  New  York,  Connecticut,  New  Jersey, 
Ottawa,  Can.,  Utah,  Arizona,  Mississippi,  Tennessee,  Alabama, 
North  Carolina,  Louisiana,  Kansas,  Iowa,  Lower  California,  Mex- 
ico, and  the  West  Indies. 

Hosts.  Wildermuth  records  this  species  from  the  following  hosts: 
Alfalfa,  cowpeas,  tomato,  almond,  Bermuda  grass,  Johnson  grass, 
wheat,  barley,  oats,  bur  clover,  yellow  sweet  clover,  soy  beans,  red 
clover,  vetch,  Hordeum  murinum,  beans,  sunflower,  cocklebur, 
Erigeron  canadensis,  mesquite,  cottonwood,  Sporobolus  airodes  and 
Trichlaris  mendocina. 

Genus  Acutalis  Fairmaire. 
In  this  genus  are  small  species  with  the  prothorax  dark  and  with 
five  apical  cells  in  the  tegmina,  the  veins  of  which  are  quite  distinct. 
A  single  species  is  reported  from  Kansas. 

Acutalis  tartafea  (Say) 

(PI.  V,  figs.  5,  6.) 

Membracis  tartarea  Say,  Jl.  Acad.  Nat.  Sci.  Phila.,  vi.  p.  242,  1830;  Compl.  Writ.,  ii, 
p.   376. 

Ceresa  tartarea  Walker,   List.  Homop.,   iv,   p.    1141,   18.52. 

Acutalis  tartarea  Uhler,  Bui.  U.  S.  Geol.  Geog.   Surv.,  i,  p.   345,   1876. 

Ceresa  semicrema  Provancher,  Pet.   Faune  Ent.   Can.,   iii,   p.    235,   1886. 

Funkhouser  gives  the  following  technical  description: 

Small  elongate  species,  very  black,  with  eyes,  undersurface  of  body  and  in 
some  cases  lateral  margins  of  pronotum  white,  apices  of  tegmina  abruptly 
hyaline. 

Head  twice  as  broad  as  long,  densely  black,  smooth,  not  punctate  nor  pu- 
bescent; eyes  prominent  and  white;  ocelli  small,  white,  about  equidistant 
from  each  other  and  from  the  eyes;  clypeus  foreshortened,  smooth,  extending 
only  slightly  in  a  semicircular  curve  below  inferior  line  of  face. 

Pronotum  intensely  black  above,  finely  punctate,  not  pubescent,  lateral 
margins  and  tip  of  posterior  process  in  some  cases  marked  with  white;  dorsal 
crest  low,  weakly  convex;  posterior  process  nearly  straight,  slightly  decurved, 
more  or  less  tectiform,  extending  beyond  abdomen  and  almost  to  end  of  apical 
cells  of  tegmina,  but  not  reaching  apex  of  hyaline  border. 

Tegmina  opaque  black  for  basal  two-thirds,  apical  third  suddenly  hyaline; 


Lawson:    Membracid^  of  Kansas.  63 

veins  heav}^  ami  black;  wide  apiral  border;   bat>al  third  i)iinctate.     Undersur- 
face  of  body  pale.    Legs  yellowish,  tarsi  fuscous. 

Length  to  apices  of  tegmina,  4.5  mm.;  width  between  humeral  angles,  2  mm. 

Internal  male  genitalia.  Styles  small,  anterior  portion  slender, 
then  wide  opposite  connective  and  narrowing  again  to  the  rather 
slender  apical  portions,  which  are  strongly  hooked,  bear  a  few  hairs, 
and  end  in  an  acute  apex;  connective  comparatively  large,  semi- 
circular; cedagus,  viewed  laterally,  quite  large,  anterior  process 
smaller  and  ending  acutely,  posterior  process  very  large  and  ending 
obtusely. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario, 
Massachusetts,  New  York,  Pennsylvania,  District  of  Columbia, 
North  Carolina,  Florida,  Mississippi,  Illinois,  Iowa,  Missouri,  Colo- 
rado, and  Utah.  Miss  Branch  reports  it  from  Douglas  county, 
Kansas. 

Hosts.    Miss  Branch  gives  Ambrosia  trifida  as  a  host. 

Genus  Micrutalis  Fowler. 

The  members  of  this  genus  are  small  and  have  but  four  apical 
cells  in  the  tegmina,  the  veins  of  which  are  very  obscure. 
A  single  member  of  the  genus  occurs  in  Kansas. 

Micrutalis  calva  (Say). 

(PI.  V,  figs.  3,  4.) 

Meinbracis  calva  Say,  Jl.  Acad.  Nat.  Sci.  Phila.,  vi.  p.  242,  1830;  Conipl.  Writ.,  ii,  p.  376. 

Membracis  melanogranuna  Perty,  Del.  An.  Art.,  pi.   35,  fig.   10,  1834. 

Smilia  flavipennis  Germar,  Silb.  Rev.  Ent.,  iii,  p.  240,  1835. 

Acutalis  flavipennis  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  497,  1846. 

Ceresa  calva  Walker,  List  Homop.,  iv,  p.   1141,   1852. 

Acutalis  melanogramma  Walker,  List  Homop.,  ii,  p.   591,   1851. 

Acutalis  calva  Fitch,  Trans.  N.  Y.  St.  Agr.  Soc,  xvi,  p.  391,  1856. 

Acutalis  illinoiensis  Coding,  Can.  Ent.,  xxv,  p.   53,   1893. 

Micrutalis  illinoiensis  Baker,  Can.  Ent.,  xxxix,  p.   116,  1907. 

Micrutalis  calva  Baker,  Can.  Ent.,  xxxix,  p.   116,   1907. 

Funkhouser  gives  the  following  technical  description: 

Very  minute;  one  of  the  smallest  species  of  Membracidae  in  the  United 
States;  usually  strongl^v  marked  with  black,  although  color  is  variable;  abdo- 
men j-ellowish;  tegmina  hyaline,  veins  very  indistinct. 

Head  broad,  smooth,  hghtly  punctate,  not  pubescent,  upper  third  black, 
lower  two-thirds  3'ellowish;  eyes  prominent,  white  or  gray;  ocelli  not  promi- 
nent, pearly,  about  equidistant  from  each  other  and  from  the  eyes  and  situated 
slightly  above  an  imaginary  line  drawn  through  centers  of  eyes;  clypeus 
rounded,  continuing  sinuate  outline  of  inferior  margin  of  face. 

Pronotum  low,  nearlj^  flat,  finelj'  punctate,  not  pubescent,  anterior  part 
usually  black,  tip  of  posterior  proce-ss  generally  pale;  posterior  process  stout, 
triangular,  just  reaching  internal  angles  of  tegmina  and  not  extending  as  far 
as  tip  of  abdomen. 


64  The  University  Science  Bulletin. 

Tegmina  entirely  hyaline,  not  punctate  nor  pubescent  at  base,  veins  indis- 
tinct, apical  border  broad.  Entire  abdomen  pale;  undersurface  of  thorax  of- 
ten marked  with  black.  Femora  black  or  ferruginous;  tibiae  fuscous,  tarsi  fer- 
ruginous. 

Length,  3-3.5  mm.;  width,  1.5-1.7  mm. 

Internal  male  genitalia.  Styles  with  anterior  and  posterior  parts 
of  about  equal  length,  anterior  ends  pointed,  strongly  swollen  op- 
posite connective,  posterior  part  curving  laterad,  with  a  distinct 
hook  before  the  acute  apex,  and  with  about  a  dozen  hairs  along 
the  lateral  margin;  connective  small,  notched  basally,  widest  at  the 
truncate  apex;  oedagus,  viewed  laterally,  U-shaped,  the  base  nearly 
straight  and  with  a  large  process  to  connective,  functional  orifice 
along  the  side  of  the  posterior  arm  which  bears  numerous  sawlike 
teeth  on  its  cephalic  aspect  and  a  few  scattered  ones  laterally. 

Distribution.  This  is  a  very  widely  distributed  species  occurring 
throughout  the  Eastern  and  Southern  states  as  shown  by  the  fol- 
lowing distribution  given  by  Van  Duzee:  New  Hampshire,  Massa- 
chusetts, New  York,  Pennsylvania,  Maryland,  District  of  Columbia, 
Virginia,  North  Carolina,  Georgia,  Florida,  Alabama,  Mississippi, 
Ohio,  Michigan,  Illinois,  Missouri,  Kansas,  Iowa,  Arkansas,  Texas, 
Colorado.     It  is  also  known  to  occur  in  the  West  Indies. 

Specimens  have  been  taken  in  the  following  Kansas  counties: 
Reno,  Douglas,  Riley,  Bourbon,  Harper,  Chautauqua,  Cowley, 
Cherokee,  Allen,  Harvey,  Montgomery,  Butler,  Kingman,  Sumner, 
Miami  and  Ottawa. 

Hosts.  Funkhouser  gives  black  locust  as  a  host.  It  is  commonly 
taken  when  sweeping  weeds  and  grasses  in  Kansas. 

Genus  Carynota  Fitch. 

The  members  of  this  genus  have  the  elytra  partially  covered  by 
the  pronotum,  which  lacks  a  horn  or  crest.  The  dorsum  is  low  and 
rounded. 

A  single  species  occurs  in  Kansas. 

Carynota  mera  (Say). 

Memhracis  viera  Say,  Jl.  Acad.  Nat.  Sci.  Phila.,  vi,  p.  301,  1831  ;   Compl.  Writ.,  ii,  p.  379. 

Carynota  mera  Fitch,  Homop.   N.  Y.  St.   Cab.,  p.   48,   1851. 

Gargara  niajus  Emmons,  Nat.  Hist.  N.  Y.  Ins.,  p.  156,  pi.   13,  fig.  6,  1854. 

Ophiderma  mera  Fitch,  Trans.  N.  Y.  St.  Agr.   Soc,  xvi,  p.   465,  1856. 

Craynota  strombergi  Goding,  Bui.  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  443,  1894. 

Funkhouser  gives  the  following  technical  description: 

Fine  large  species;  gray  marked  with  dark  brown  and  chestnut;  pronotum 
convex  and  elevated;  tegmina  fuscous-hyaline  tipped  with  dark  brown. 

Head  nearly  twice  as  broad  as  long,  uniform  light  gray,  very  distinctly 
punctate,  sparingly  pubescent  with  short,  white  hairs;   eyes  very  prominent 


Lawson:    Membracid^  of  Kansas.  65 

and  lirowu;  ocrlli  proiniiunt,  pc.-iiiy.  iiiai]LiiiuMl  wil  li  i)range.  somewhat  pro- 
tniding,  nearer  to  each  otlier  tliaii  to  the  eyes;  clypeus  subtriangular,  con- 
tinuing inferior  outline  of  fac(\  tip  i>roduced  in  small  tooth,  hirsute. 

Pronotvun  gray,  finely  punctate.  ]>ubescent.  median  carina  percurrent ;  meto- 
pidium  convex,  irregular  brown  mark  above  internal  angle  of  each  eye;  dorsal 
line  arcuate,  suddenly  depressed  before  posterior  process  in  female,  depression 
not  so  evident  in  male;  wide,  dark  brown,  transverse  band  crossing  middle  of 
pronotum  on  each  side;   jiosterior  jirocess  heavy,  pointed,  tip  chestnut. 

Tegmina  smoky  hyaline,  veins  prominent,  bases  punctate,  especially  along 
veins  and  at  costal  margins,  tips  dark  brown  or  black.  Legs  and  undersurface 
of  body  ferruginous. 

Length:  female,  10  mm.;  male,  8.5  mm.  Width:  female,  5  mm.;  male 
4  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York,  New  Jersey,  Pennsylvania,  North  Carolina,  Ohio,  Illinois, 
Iowa,  Missouri  and  Texas.  The  writer  took  a  single  specimen  in 
Ottawa  county,  Kansas. 

Hosts.  Funkhouser  mentions  hickory,  butternut,  pecan  and  oak 
as  hosts  of  this  species. 

Genus  Thelia  Amyot  &  Serville. 

The  members  of  this  genus  are  at  once  recognized  by  the  pos- 
session of  a  long  horn  on  the  anterior  part  of  the  prothorax,  which 
points  upwards  and  forwards. 

Both  members  of  the  genus  known  to  occur  in  the  United  States 
are  found  in  Kansas.    They  may  be  separated  by  the  following  key : 

KEY   TO   SPECIES. 

A.    Prothorax  of  male  without  lateral  yellow  stripe.  uhleri. 

AA.   Prothorax  of  male  with  lateral  yellow  stripe.  bimaculata. 

Thelia  uhleri  Stal. 

Thelia   uhleri  .Stal,   Of.   Vet.   Akad.    Forh.,   xxvi,   p.    248,    18G9. 

The  original  description  follows: 

Griseo-ferruginea,  pilosa,  thorace  remote  pallido-consperso ;  tegminibus 
sordide  hyalinis,  apice  fusco-nebulosis,  basin  versus  punctatis.  Female:  Long. 
9;  cum  cornu,  13.    Lat.  4%  mill.     Wisconsin.     (Mus.  Holm.) 

T.  himaculatce  maxime  affinis,  pictura  thoracis,  ejusdem  cornu  antico 
paullo  longiore  processuque  postico  rugis  longitudinalibus  destituto  divergit. 
Caput  remote  punctulatum.  Thorax  dense  distinctque  punctatus,  angulis 
lateralibus  nonnihil  prominulis.  rectis,  cornu  antico  antrorsum  valde  nutante, 
processu  postico  apicem  tegminum  subattingent(\ 

Ad  hanc  speciem  verisimiliter  spectat  Fitch,  quiim  dicit  Membracem  belli- 
geram  Say  ad  Theliam  referendam  esse;  hsec  species  autem  ad  Platycotem 
pertinet  et  eadem  est  ac  P.  sagittata  Germ.,  quae  American!  borealem  (nee 
Brasiliam)  inhabitat. 


66  The  University  Science  Bulletin. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York,  Pennsylvania,  Michigan,  Illinois,  Wisconsin,  and  Kansas. 

Hosts.   Seemingly  unknown. 

The  writer  has  not  seen  specimens  of  this  species,  but  it  is  included 
in  this  paper  because  of  Van  Duzee's  records. 

Thelia  bimaculata  (Fabricius). 

(PI.   V,   figs.   1,   2.) 

Membracis   bimaculata  Fabricius,   Ent.   Syst.,   iv,    p.    10,   1794. 
Hemiptycha  blnotata  Harris,  Kept.  Ins.  Mass.,  p.   179,   1841. 
Hemiptycha  acuviinata  Harris,   Rept.   Ins.   Mass.,  p.    179,    1841. 
Thelia  biinaculata  Amyot  &  Serville,   Hemip.,   p.   541,    1843. 
Thelia  unanimis  Walker,  List  Homop.,  ii,  p.   566,   1851. 

Funkhouser  gives  the  following  technical  description: 

Female.  Gray  with  indistinct  darker  irregular  markings;  porrect  cylindrical 
horn  slightly  flattened  and  somewhat  darker  in  color  at  tip;  tegmina  hyaline, 
apices  fuscous,  almost  reaching  extremity  of  dorsal  process. 

Head,  including  eyes,  twice  as  broad  as  long,  grayish-yellow  mottled  with 
ferruginous  and  brown;  margins  of  lorae  strongly  sinuate;  eyes  dark  brown; 
ocelli  white,  nearer  to  each  other  than  to  the  eyes  and  situated  on  a  line  drawn 
through  centers  of  eyes;  clypeus  pilose;  beak  extending  to  posterior  coxae; 
head  very  sparingly  punctate  and  sparsely  pilose. 

Thorax  gray,  deeply  and  densely  punctate;  median  percurrent  brown  line 
sharpened  into  a  ridge  on  extremity  of  horn  and  at  apex  of  posterior  process; 
sides  of  prothorax  roughly  and  irregularly  carinate;  horn  porrect  and  greatly 
variable  in  length;  cylindrical  except  at  extreme  tip,  where  it  is  flattened 
laterally;  posterior  process  heavy,  tectiform,  gradually  acute,  almost  straight, 
very  slightly  decurved  and  extending  beyond  apices  of  tegmina. 

Tegmina  hyaline,  apices  fuscous,  bases  and  costal  regions  lightly'  punctate; 
underwings  hyaline,  two-thirds  as  long  as  tegmina.  Undersurface  of  body 
gray-brown,  pubescent.  Legs  uniform  yellow-brown;  femora  thick  and 
smooth;  tibiae  and  tarsi  densely  pilose. 

Length,  11  mm.,  including  horn,  14  mm.;  width  between  humeral  angles, 
5.5  mm. 

Male:  Differs  from  female  in  size  and'  markings.  Smaller,  body  some- 
what less  robust;  porrect  horn  usually  shorter  and  tending  to  curve;  tegmina 
equalling  apex  of  posterior  process.  Color  deep  chocolate  brown;  porrect 
horn  almost  black;  apex  of  posterior  process  becoming  cinnamon  brown;  a 
wide,  brilliant,  lemon-yellow  longitudinal  stripe  on  each  side  of  prothorax, 
extending  from  margin  halfway  to  median  dorsal  line,  also  small  patches  of 
yellow  on  metopidium;  head  yellow  with  brown  patches.  Undersurface  of 
abdomen  darker  than  in  female. 

Internal  male  genitalia.  Styles  with  anterior  portion  short, 
posterior  part  longer,  wide,  and  parallel-margined  till  near  apices, 
which  are  curved  strongly  laterad  and  end  truncately  but  with  a 
distinct  recurved  hook,  the  curved  apices  bearing  scattered  short 
hairs;  connective  large,  rather  distinctly  seven-sided;  oedagus,  viewed 
laterally,  with  a  slender  process  to  anal  tube,  the  apical  portion 


Lawson:    Mkmbracid.e  of  Kansas.  67 

very  large,  club-f^haped,  tlie  runctional  orilice  just  before  the  ex- 
treme apex,  the  anterior  aspeet  of  the  apex  with  many  filelike  teeth. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario, 
Massachusetts,  New  York,  New  Jersey,  Pennsylvania,  North  Car- 
olina, Ohio  and  Illinois.  The  writer  has  taken  it  in  Douglas  county, 
Kansas. 

Hosts.    Black  locust  seems  to  be  the  only  host. 

Genus  Glossonotus  Butler. 
The  members  of  this  genus  possess  a  tongue-shaped  crest,  which 
arises  from  between  the  humeral  angles. 

A  single  species  of  the  genus  is  recorded  from  Kansas. 

Glossonotus  acuminatus  (Fabricius). 

Membracis  acuminata  Fabricius,  Syst.  Ent.,  P.  675,  1775. 

Cicada  acuminata  Ginelin  in  Linnaeus,  Syst.   Nat.,  edn.    13.  i,   pt.   4,   2094,  1778. 

Centrotus  acuminata  Fabricius,  Syst.   Rliyng.,  p.    18,   1803. 

Thelia  acuminata  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  310,  pi.   5,  fig.  15,  1846. 

Hemiptycha  acuminata  Harris,  Treat.  Ins.   Inj.   A'eg.,  edn.   3,   p.   223,   1862. 

Telamona  acuminatus  Stal,  Hemip.  Fabr.,  ii,  p.  115,  1869. 

Glossonotus  acuminata  Butler,  Cist.  Ent.,  ii,  p.  222,  1877. 

Thelia  crataegi  Smith,  Ins.  N.  J.,  p.  441,  1890. 

Funkliouser  gives  the  following  technical  description: 

Dark  gray  mottled  with  brown;  dorsal  crest  high,  flattened  and  swollen  at 
tip;  humeral  angles  prominent  and  triangular;  tegmina  hyaline  tipped  with 
brown,  veins  punctured. 

Head  almost  as  long  as  wide,  gray  with  distinct  scattered  black  punctures 
and  fine  whitish  pubescence;  base  sinuate;  eyes  large,  prominent,  brown, 
extending  as  far  as  bases  of  humeral  angles;  ocelli  large,  prominent,  pearly 
with  white  margins,  nearer  to  each  other  than  to  the  eyes;  clypeus  continuing 
inferior  line  of  face,  punctate  with  black,  pubescent  with  white,  tip  prolonged 
into  a  point;  antennae  long  and  well  developed. 

Pronotum  dark  gray  with  irregular  markings  of  brown,  coarsely  and  regu- 
larh'  punctate  with  black,  very  sparingly  pubescent;  metopidium  convex, 
median  carina  prominent  and  decorated  with  alternate  lines  of  brown  and 
yellowish,  irregular  black  markings  above  internal  angles  of  eyes,  humeral 
angles  prominent,  triangular,  flattened,  acute;  pronotal  crest  almost  as  high 
as  length  of  pronotum,  widened  and  flattened  at  tip,  margin  decorated  with 
pale  areas,- projecting  usually  forward  as  well  as  upward;  posterior  process 
gradually  acuminate,  reaching  apices  of  tegmina. 

Tegmina  hyaline,  tips  clouded  with  smoky  brown,  bases  and  margins  of 
veins  punctate,  veins  prominent.  Undersurface  of  thorax  fuscous;  abdomen 
ferruginous.    Legs  fuscous  marked  with  brown. 

Length.  10  mm.;  width  between  tips  of  humeral  angles,  6  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  Mas- 
sachusetts, New  York,  New  Jersey,  Pennsylvania,  Michigan,  Iowa, 
Kansas  and  Arkansas. 

Hosts.    Funkhouser  gives  young  white  oak  as  the  host. 


68  The  University  Science  Bulletin. 

Genus  Heliria  Stal. 

The  members  of  this  genus  are  characterized  by  having  a  dis- 
tinctly step-shaped  dorsal  crest. 

Two  members  of  the  genus  are  known  to  occur  in  Kansas.  These 
Van  Duzee  separates  by  the  following  key : 

KEY   TO   SPECIES. 

A.  Larger,  12-13  mm.;  grey  or  brownish  grey;  posterior  foliole  of  the  crest 
Httle  elevated  above  the  dorsal  line  anteriorly,  its  hind  angle  subacute; 
humeral  angles  greatly  produced.  cristata. 

AA.    Smaller,  8  mm.,  darker  brown;  posterior  foliole  of  the  crest,  when  well 
differentiated,  elevated  at  least  its  own  width  above  the  dorsal  line. 

scalaris. 
Heliria  cristata   (Fairmaire). 

Thelia  cristata  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  311,  pi.   5,  fig.  14,  184fi. 
Telavxona  jagi  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  51,  1851. 

Telamona  acclivata  Emmons,  Nat.  Hist.  N.  Y.  Ins.,  p.   155,  pi.  3,  fig.  5,  1854. 
Heliria  cristata  Stal,  Of.  Vet.  Akad.  Forh.,  xxiv,  p.  556,  1867;   .xxvi,  p.   249,  1869. 
Telamona  cristata  Fowler,  Biol.  Centr.  Am.,  Homop.,  ii,  p.  144,  pi.  9,  fig.  6,  1896. 

The  following  is  the  original  description : 

T.  cristata.    Mexique.    Long.  0.012. 

Praecedente  differt  testaceo-obscuro  et  tuberculorem  dispositione ;  primo 
antice  inclinato,  secundo  sat  fortiter  acuto. 

Xe  differe  de  I'espece  precedente  que  par  se  couleur  d'un  testace  obscue, 
et  la  disposition  des  lobes  dorsaux  dont  I'anterieur  est  incline  en  avant,  et  le 
second  plus  aigu.    Coll.  Signoret. 

The  preceding  species  referred  to  above  is  H.  scalaris. 

Distribution.  Van  Duzee  reports  this  species  from  New  York, 
New  Jersey,  North  Carolina,  and  Illinois.  Popenoe  reports  it  from 
Kansas. 

Hosts.    Unknown. 

Heliria  scalaris  (Fairmaire). 

Thelia  scalaris  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  311,  1846. 

Telamona  jagi  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  51,  1851. 

Thelia  scalaris  Walker,  List  Homop.,  ii,  p.  565,  1851. 

Heliria  scalaris  Stal,  Of.  Vet.  Akad.  Forh.,  xxiv,  p.  556,  1867;   xxvi,  p.  249,  1869. 

Telamona  scalaris  Butler,  Cist.  Ent.,  ii,  p.  222,  1877. 

Funkhouser  gives  the  following  technical  description: 

A  small  species,  uniform  brown  in  color;  crest  as  high  as  its  length  at  base;' 
posterior  process  not  reaching  apices  of  tegmina;  tegmina  smoky  hyaline,  tips 
browTi. 

Head  as  wide  as  long,  sculptiu-ed.  yellowish,  irregularly  punctuate  with 
brown,  sparingly  pubescent;  base  strong^  sinuate;  eyes  prominent,  brown, 
reaching  base  of  humeral  angles;  ocelli  prominent,  translucent,  nearer  to  each 
other  than  to  the  eyes;  clypeus  extending  below  inferior  margin  of  face, 
j'ellowish,  punctin-ed  with  brown,  pubescent. 

Pronotum  imiform  brown,  coarsely  punctured;  dorsal  crest  swollen  at  base, 
flattened  at  apex,  as  high  as  its  length  at  base,  distinctly  step-shaped,  anterior 


Lawson:    Membracid.e  of  Kansas.  69 

lobe  rouiultHl  ami  inDjei'tin^  I'orwanl,  i)Ot>tciior  lobe  sharply  angular,  two-thirds 
as  hifih  as  anterior,  both  lobes  in  some  cases  margined  with  patches  of  darker 
brown;  posterior  process  short,  heavy,  acute,  not  reaching  apices  of  tegmina; 
humeral  angles  triangular,  flattened,  blunt. 

Tegmina  smoky  hyaline,  bases  dark  brown  and  punctate,  tijis  brown,  veins 
heavy  and  often  ])unctured  along  margins,  l^ndersurface  of  thorax  ferruginous, 
segments  margined  with  paler;  abdomen  brown.  Legs  ferruginous;  tibiae  and 
tai-si  hairy. 

Length,  8  mm.;  width,  4.8  mm. 

Distribution.  Van  Diizec  reports  this  species  from  Ontario,  New 
York,  Ne\v  Jersey,  Pennsylvania,  Illinois  and  Colorado.  There  is  a 
specimen  in  the  Snow  collection  from  Kansas  City,  Mo.,  so  it  un- 
doubtedly occurs  in  eastern  Kansas. 

Hosts.    Unknown. 

Genus  Telamona  Fitch. 

The  following  is  the  original  description  of  the  genus: 

Humeral  angles  projecting,  pointed  and  earlike:  dorsum  compressed-folia 
ceous,  the  keel  abruptly  elevated  at  one  or  both  its  ends,  forming  a  somewhat 
square  crest  or  foliole;  thorax  nearly  or  quite  reaching  the  tips  of  the  elytra, 
with  elevated  longitudinal  lines  on  each  side;  apical  cellule  triangular,  its  end 
rounded.  The  squarish  dorsal  crest  forms  a  marked  distinction  between  the 
genus  here  proposed,  and  that  of  Thelia,  to  which  it  is  mo&l  nearly  related. 

Eleven  members  of  this  genus  are  known  to  occur  in  Kansas. 
These  may  be  separated  by  the  following  key : 

KEY   TO   SPECIES. 

A.    Anterior  margin  of  crest  sloping  to  front  of  metopidium,  without  obvioua 
sinus  at  its  anterior  base. 
B.    Crest,  if  elevated,  more  or  less  rounded  or  pointed  at  apex. 
C.    Crest  distinctly  elevated. 

D.   Brownish  species;  crest  higher  and  triangular. 

pyramidata. 
DD.   Greenish  species;  crest  an  obtusely  conical  hump. 

viridia. 
CC.    Crest  scarcely  elevated,  broadly  rounded  above.      obsoleta. 
BB.   Crest  elevated,  square  or  nearly  so  at  apex. 
C.   Crest  not  as  high  as  broad. 

D.    Greenish-brown  species.  lugubris. 

DD.    Gray,  with  oblique  brown  fascia.  decorata. 

CC.    Crest  as  high  or  higher  than  broad.  querci. 

A  A.    Crest  nearly  vertical  before  or  sometimes  overhanging. 
B.    Color  greenish  or  j^ellowish. 

C.    Females  bright  green;  males  yellow,  banded  with  brown. 

unicolor. 
CC.   Both  sexes  of  about  the  same  color. 

D.    Pronotal  hump  very  high.  extrema. 

DD.    Pronotal  hump  moderate.  collina. 


70  The  University  Science  Bulletin. 

BB.   Color  not  green. 

C.   Yellow,  mottled  with  brown.  tristis. 

CC.    Gray,  with  transverse  brown  band.  ampelopsidis. 

Telamona  pyramidata  Uhler. 

(PI.  V,  figs.  7,  8.) 

Telamona  pyramidata  Uhler,  Wheeler's  Rept.  Chief  Eng.   for  1877,  p.  1333. 
Telonaca  pyramidata  Ball,  Proc.  Biol.  See.  Wash,  xxxi,  p.  28,   1918. 

Funkhouser  gives  the  following  technical  description: 

Long,  narrow  body;  crest  triangular  and  pyramidal,  as  the  name  would 
suggest;  mottled  brown  with  a  dark  transverse  fascia  extending  from  tip  of 
crest  to  lateral  margin  of  pronotum,  and  a  second  shorter  fascia  behind  it; 
posterior  process  extending  to  tips  of  tegmina;  tegmina  hyaline,  punctate  at 
bases,  brown  at  apices.  Differs  from  T.  declivata  chiefly  in  shape  of  dorsal 
crest. 

Head  wider  than  long,  yellowish  with  large  irregular  punctures  of  brown, 
sparingly  pubescent;  base  regularly  sinuate;  eyes  large,  prominent,  gray;  ocelli 
large,  prominent,  somewhat  protruding,  translucent;  clypeus  subtriangular, 
sutures  distinct,  apex  slightly  produced,  hairy. 

Pronotum  deeply  punctate,  not  pubescent;  metopidium  convex,  decorated 
with  patches  of  yellowish  and  dark  brown,  median  carina  prominent,  heavy, 
black,  broken  by  circular  areas  of  yellowish ;  humeral  angles  prominent,  tecti- 
form,  blunt,  brownish;  dorsal  crest  triangular,  rounded  at  tip,  margin  flattened 
and  brown,  posterior  margin  pale ;  posterior  process  long,  slender,  slightly  curv- 
ing downward,  extending  beyond  tips  of  tegmina;  median  carina  percurrent. 

Tegmina  hyaline,  bases  and  costal  margins  coarsely  punctate  but  not  pu- 
bescent, tips  brown.  Undersurface  of  thorax  flavous;  abdomen  dark  brown. 
Legs  yellowish ;  tibiae  mottled  with  brown,  hairy ;  tarsi  flavous ;  claws  fer- 
ruginous. 

Length,  9-11  mm.;  width,  5-6  mm. 

Internal  male  genitalia.  Styles  with  anterior  portion  broad  and 
flat,  posteriorly  curved  outward  and  with  a  large,  blunt  apical  hook, 
the  apical  fourth  with  a  few  short  hairs ;  connective  large  and  heart- 
shaped,  bearing  three  longitudinal  ridges;  oedagus,  viewed  laterally, 
U-shaped,  anterior  process  rather  slender  and  with  a  distinct  tooth 
on  its  cephalic  aspect,  posterior  process  large  and  club-shaped,  the 
apex  with  sawlike  teeth  on  its  anterior  surface,  and  the  functional 
orifice  on  the  posterior. 

Distribution.  Gillette  and  Baker  report  this  species  from  Colo- 
rado and  Funkhouser  from  New  York.  The  writer  has  taken  it  at 
Ames,  Iowa.  Specimens  have  been  taken  in  Kansas  from  Johnson, 
Pawnee,  Cherokee,  Riley  and  Pottawatomie  counties.  It  seemingly 
occurs  throughout  the  Eastern  states  and  west  to  the  Rocky  Moun- 
tains. 

Hosts.    Funkhouser  reports  it  on  chestnut  oak.    Gillett  and  Baker 


Lawson:    Membracid.e  of  Kansas.  71 

give  Cottonwood  and  Virginia  creeper.     The  writer  has  taken  it 
abundantly  on  willow. 

Tlie  writer  has  been  unable  to  find  any  characters  that  would 
seem  to  separate  this  species  from  the  members  of  the  genus  Tela- 
mona.  The  genitalia  are  precisely  as  in  other  members  of  the 
genus.  Hence  it  is  our  belief  that  the  genus  Telonaca  should  be 
sunk  as  a  synonym  of  Telamona. 

Telamona  viridia  Ball. 

(PI.  VII,  figs.  7,  8.) 
Telamona  viridia  Ball,  Proc.  Biol.  Soc.  Wash.,  xvi,  p.  178,  pi.  1,  fig.  3,  1903. 

The  original  description  follows: 

Resembling  pyramidata  in  size  and  form,  but  with  less  of  a  hump.  Grass- 
green,  the  male  with  some  fuscous  on  posterior  half  of  hump  and  again  at 
apex  of  pronotum. 

Length:   female,  11  mm.;  male,  9  mm.    Width,  female,  5.3  mm. 

Pronotal  hump  in  the  shape  of  an  obtuse  pyramid  one-third  the  distance 
back  from  eye  to  apex  of  pronotum,  a  slight  angle  on  posterior  margin  just 
below  apex  especially  marked  in  the  male.  Height  of  hump  slightly  less  than 
one-third  the  pronotal  length.  Humeral  angles  broad,  slightly  rounded,  a 
trifle  longer  than  eye. 

Color.  Female,  grass-green  slightly  mottled  with  yellow;  carina  light  ex- 
cept at  apex  of  hump  and  at  tip,  where  it  is  tawny.  Male,  grass-green;  carina 
light  interrupted  with  tawny;  a  fuscous  band  nans  obliquely  backward  from 
apex  of  hump  and  fades  out  before  reaching  the  pronotum  proper,  or  some- 
times connects  with  a  tawny  spot  on  lower  margin,  whole  apex  of  pronotum 
tawny. 

Internal  male  genitalia.  Styles  large,  anterior  end  flattened,  pos- 
terior part  stout,  curved  strongly  laterad  apically,  bluntly  hooked, 
and  bearing  a  few  scattered  hairs;  connective  large,  heart-shaped, 
with  three  longitudinal  ridges;  oedagus,  view^ed  laterally,  U-shaped, 
anterior  arm  more  lightly  chitinized  and  with  short  blunt  anterior 
tooth,  posterior  arm  club-shaped,  the  apex  anteriorly  with  many 
small  filelike  teeth,  posteriorly  with  the  functional  orifice. 

Distribution.  Ball  reports  the  type  specimens  from  Colorado, 
Iowa  and  Illinois.  There  is  a  specimen  in  the  Snow  collection,  taken 
by  F.  X.  Williams,  from  Gove  county,  and  a  single  specimen  has 
been  taken  in  Riley  county,  Kansas. 

Hosts.    Ball  gives  cottonwood  {Populus  monilifera)  as  the  host. 

Telamona  obsoleta  Ball. 

Telamona  obsoleta  Ball,  Proc.  Biol.   Soc.  Wash.,  xvi,  p.  178,  pi.   1,  fig.   2,  1903. 

The  following  is  the  original  description: 

Resembling  irrorata,  but  smaller  and  with  a  smaller  and  more  rounding 
hump.    Length:    female,  10  mm.;  male,  9  mm.    Width,  5  mm. 


72  The  University  Science  Bulletin. 

Dorsal  hump  low  and  much  inflated;  it  scarcely  narrows  from  the  base  to 
just  before  the  apex  where  it  rounds  in  to  form  a  carina.  Anterior  margin 
rising  just  back  of  the  humeral  angles  and  extending  from  there  half  way  to 
the  apex  of  the  pronotum.  The  height  is  about  equal  to  the  whole  length 
and  it  rounds  down  to  the  pronotum  proper  at  both  extremities.  Front  much 
elevated  above  the  level  of  the  eyes  so  that  the  ocelli  are  farther  from  the 
base  of  front  than  from  each  other. 

Color.  Yellow  with  the  punctures  fuscous,  sometimes  coalescing  into  brown- 
ish fuscous  spots,  giving  the  whole  insect  an  irrorate  and  mottled  appearance, 
with  little  regularity  of  pattern.  Usually  there  is  a  semicircle  of  lighter  shade 
back  of  the  humeral  angles  and  a  light  spot  on  middle  of  hump.  There  is  a 
pair  of  large,  straggling  black  marks  above  and  within  the  eyes,  some  brown 
on  the  inner  nervures  of  corium,  and  a  smoky  brown  cloud  at  apex. 

Distribution.  Ball  reports  this  species  from  Iowa  and  from  Kan- 
sas. Van  Duzee  records  it  from  Illinois  and  Fmikhouser  from  New 
York.  In  Kansas  it  has  been  taken  in  Pottawatomie  and  Mont- 
gomery counties. 

Hosts.    Ball  gives  elm  as  a  host;  Funkhouser  reports  it  from  oak. 

Telamona  lugubris  Ball. 

Telamona  lugubris  Ball,  Proc.  Biol.  Soc.  Wash.,  xvi,  p.   179,   1903. 

The  original  description  follows: 

Form  of  reclivata  nearly,  slightly  shorter  and  stouter  built  and  with  a  lower 
and  longer  hump  and  lacking  the  markings  of  that  species.  Obscurely  greenish 
brown.    Length:    female,  11  mm.;  male,  9.5  mm.    Width,  5.5  mm. 

Dorsal  hump  of  moderate  size,  arising  just  back  of  lateral  angles;  anterior 
margin  sloping  back,  forming  a  right  angle  with  the  inclined  crest,  posterior 
margin  perpendicular  or  slightly  overhanging.  Base  of  hump  occupying  a 
little  over  two-fifths  of  distance  from  humeral  angles  to  apex  of  pronotum. 
Humeral  angles  blunt  and  obtuse,  about  two-thirds  as  long  as  the  eye. 

Color.  Pale  yellow,  the  more  or  less  darkened  punctures  giving  the  insect 
a  general  grayish  cast  with  still  darker  shadings  on  the  lateral  faces  of  the 
hump  and  sometimes  on  the  apex  of  pronotum. 

Distribution.  Ball  reports  this  species  from  Iowa  and  also  from 
Pottawatomie  county,  Kansas.  Doctor  Funkhouser  has  specimens 
from  New  Mexico. 

Hosts.    Ball  gives  scrub  oak  as  the  host. 

Telamona  decorata  Ball. 

Telamona  decorata  Ball,  Proc.  Biol.  Soc.   Wash.,  xvi,  p.   179,   1903. 

The  original  description  follows: 

Smaller  than  lugubris,  with  a  shorter  and  more  rounding  hump.  Yellowish 
fuscous  with  the  hump  deep  testaceous  brown.    Length,  9  mm.;  width,  4.5  mm. 

Dorsal  hump  sloping  uj)  from  both  front  and  rear,  crest  rounding,  highest 
just  in  front  of  the  middle,  hump  occupying  scarcely  two-fifths  of  the  pronotum 
from  the  humeral  angles  back.  Humeral  angles  short  and  blunt,  about  tw)- 
thirds  the  length  of  the  eye. 


Lawsox:    Membracid^  of  Kansas.  73 

Color.  Face  and  pronotiini  pale  yellow,  very  slightly  washed  with  brown 
in  the  female,  and  with  a  definite  brown  shade  in  the  male;  median  carina 
alternately  light  and  dark  l^efore  the  iiuni)).  Hiimii  rieli  testaceous  with  a  few 
light  spots  on  the  sides,  a  definite  light  mark  at  the  base  in  front,  which  may 
extend  up  on  to  the  carina,  and  the  whole  posterior  margin  light.  This  latter 
light  spot  extends  down  on  to  the  pronotum  and  connects  with  an  irregular 
transverse  light  band  about  halfway  to  apex.  Each  side  of  this  band  is  an 
irregular  testaceous  band,  the  anterior  one  connected  with  the  testaceous 
hump.    A  pair  of  spots  above  the  eyes  and  the  apex  of  elytra  brownish  fuscous. 

Distnbution.  Ball  described  this  species  from  specimens  taken 
in  Iowa,  Arkansas,  and  Pottawatomie  county,  Kansas.  Fimkhouser 
reports  it  from  New  York. 

Hosts.    Ball  mentions  red  oak  as  a  host. 

Telamona  querci  Fitch. 

Telamona  querci  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  51,  1851. 

Telamona  quercus  Walker,  List  Homop.,  iv,  p.  1145,  1852. 

TheUa  quercus  Smith,  Cat.  Ins.  N.  J.,  edn.  1,  p.  441,  1891. 

Telamona  brunneipcntiis  Buckton,  Monog.  Membr.,  p.  197,  pi.  43,  fig.   1,   1903. 

Funkhouscr  gives  the  following  technical  description: 

Very  close  to  T.  monticola;  pronotum  shorter,  darker;  dorsal  crest  with 
prominent  pale  fascia  on  posterior  margin;  tegmina  nearh'  hyaline,  tips 
faintly  clouded. 

Head  roughly  sculptured,  flavous  mottled  with  brown,  faintly  longitudinally 
striate,  ^■erJ'  faintly  punctate,  pubescent,  base  weakly  sinuate;  eyes  prominent, 
dark  bro^Mi;  ocelli  very  prominent,  protruding,  brownish,  margins  pale,  much 
nearer  to  each  other  than  to  the  eyes ;  clypeus  nearly  flat,  punctate,  pubescent, 
base  marked  with  brown,  tip  extended  below  inferior  margin  of  face. 

Pronotum  densely  but  finely  punctate,  sparingly  pubescent,  dark  brown 
mottled  with  green;  metopidium  convex,  median  carina  prominent,  black  in- 
terrupted with  pale  green ;  humeral  angles  short  and  blunt ;  dorsal  crest  sloping 
backward,  longer  than  high,  higher  before  than  behind,  posterior  margin  dis- 
tinctly pale;  posterior  process  short,  acute,  marked  with  greenish  before  apex, 
not  reaching  tips  of  tegmina. 

Tegmina  hyaline,  bases  punctured  but  not  pubescent,  tips  clouded  with 
brown,  veins  brown.    Undersurface  of  body  brown.    Legs  flavous;  tibiae  haiiy. 

Length  of  pronotum,  9  mm.;  to  tips  of  tegmina,  11  mm.;  width,  5.5  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York.  District  of  Columbia,  Ohio,  Colorado,  New  Mexico,  and  Ne- 
vada. Popenoe  reports  this  species  from  Kansas,  and  there  is  a 
specimen  in  the  Snow  collection  from  Kansas  City,  Mo.,  so  that  it 
doubtless  occurs  in  the  state.  The  writer  took  a  single  specimen  of 
the  species  at  St.  Paul,  Minn. 

Hosts.    Funkhouser  gives  white  and  chestnut  oak  as  hosts. 


74  The  University  Science  Bulletin. 

Telamona  unicolor  Fitch. 

Telamona  unicolor  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  50,  1851. 
Telamona  fasciata  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  50,  1851. 
Hemiptycha  diffusa  Walker,  List  Homop.,  Suppl.,  p.  143,  1858. 

Funkhouser  gives  the  following  technical  description: 

Females  large,  brilliant  iirufonn  grass-green;  males  smaller,  bright  yellow 
with  deep  brown  fascia.  Very  striking  in  color;  large  size;  crest  high  and 
square;  tegmina  tipped  with  brown. 

Female.  Head  nearly  twice  as  wide  as  long,  green  punctate  with  brown, 
finely  pubescent;  eyes  large,  brown;  ocelli  large,  orange,  nearer  to  each  other 
than  to  the  eyes;  clypeus  deeply  punctate,  pubescent,  tip  in  a  pointed  ex- 
tension. 

Pronotum  concolorous  green,  fading  to  mottled  yellow  in  cabinet  specimens ; 
very  finely  punctate  and  pubescent;  metopidium  more  or  less  angular,  median 
carina  distinct,  three  small  brown  spots  mesad  of  humeral  angles;  humeral 
angles  produced,  triangular,  blunt;  crest  large,  high,  much  higher  before  than 
behind,  anterior  margin  less  sloping  than  posterior,  dorsal  margin  brownish; 
posterior  process  long,  gradually  acute,  apex  brownish  and  not  reaching  tips 
of  tegmina. 

Tegmina  brownish  hyaline,  bases  and  costal  regions  punctate  with  black, 
tips  clouded  with  dark  brown,  veins  prominent.  Undersurface  of  thorax 
flavous,  abdomen  yellowish,  pubescent,  ovipositor  brown.  Legs  flavous;  tibiae 
mottled  with  brown;  tarsi  ferruginous. 

Length,  11  mm.;  width,  6  mm. 

Male.  Differs  from  female  in  size  and  color.  Head  mottled  brown  and  yel- 
low, much  darker  than  that  of  female,  much  sculptured,  inferior  line  of  face 
strongly  sinuate. 

Pronotum  bright  yellow,  metopidium  strongly  shaded  with  brown;  dark 
brown  fascia  on  front  of  dorsal  crest;  dark  brown  fascia  on  posterior  third  of 
crest  extending  gradually  narrowed  to  lateral  margin  of  pronotum;  posterior 
median  line  of  crest  yellow,  transverse  band  of  yellow  behind  crest;  apex  of 
posterior  process  brown. 

LTndersurface  of  body  deep  brown.  Legs  strongly  fiavous  marked  with 
brown. 

Length,  10  mm.;  width,  5  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York,  New  Jersey,  Pennsylvania,  North  Carolina,  Michigan,  Il- 
linois, Iowa,  Kansas,  Missouri  and  Texas.  In  Kansas  it  has  been 
taken  in  Pottawatomie  county. 

Hosts.  Funkhouser  gives  hickory,  butternut,  walnut  and  bass- 
wood  as  hosts. 

Telamona  extrema  Ball. 

Telamona  extrema  Ball,  Proc.  Biol.  Soc.  Wash.,   xvi,  p.    179  pi.   1,  fig.   1,   1903. 

The  original  description  follows : 

Form  of  unicolor  nearly,  smaller,  and  with  a  still  longer  hump.  Greenish 
testaceous.    Length:    female,  10  mm.;  male,  9  mm.    Width,  5  mm. 


Lawson:    Membracid.e  of  Kansas.  75 

Pronotal  hump  vpry  high,  almost  quadrate,  occupying  the  anterior  three- 
fifths  of  pronotum,  anterior  margin  rising  perpendicularly  from  face,  crest 
highest  just  back  of  the  well  rounded  anterior  angle  from  which  it  slopes 
slightly  to  the  almost  perpendicular  posterior  face.  Humeral  angles  moderate, 
as  long  as  the  eyes. 

Color:  Greenish  testaceous;  a  spot  above  each  eye  and  the  median  carina 
back  to  the  posterior  angle  of  hump  fuscous;  posterior  face  of  hump  broadly 
marked  with  creamy  white,  which  narrows  to  a  hne  on  the  carina  posteriorly 
in  the  female,  and  disappeai-s  entirely  in  the  male.  The  lower  margin  of  the 
humeral  angles  is  sometimes  marked  with  fuscous. 

Distribution.  This  species  was  described  from  specimens  taken  in 
Iowa,  and  Marion  comity,  Kansas.  Van  Duzee  reports  it  also  from 
Massachusetts,  Rhode  Island  and  New  Jersey. 

Hosts.    Ball  gives  oak  as  a  host. 

Telamona  collina  (Walker). 

Thelia  collina  Walker,  List  Homop.,  ii,  p.  565,   1851. 

Telamona  collina  Butler,  Cist.     Ent.,  ii,  p.   220,   1877. 

Telamona  turritella  Buckton,  Monog.  Membr.,  p.  198,  pi.  44,  fig.  6,  1903. 

The  following  is  the  original  description: 

Testacea  vel  viridi-flava;  prothorax  ai)ice  niger;  carina  ferruginea  vel  fulva; 
alse  limpidse;  alae  anticte  basi  fulv». 

Testaceous,  shining:  head  finely  punctured,  short,  almost  transversely 
spindle-shaped,  much  narrower  than  the  fore-chest,  impressed  on  each  side  of 
the  disk,  with  five  slight  undulations  along  the  hind  border,  and  seven  on  the 
fore  border;  a  slight  fuiTow  extends  from  the  hind  border  to  the  face,  whose 
hind  side  is  slightly  obconical  and  occupies  less  than  half  the  length  of  the 
face:  fore-chest  roughly  punctured,  convex  and  with  a  slight  middle  ridge 
in  front,  forming  on  each  shoulder  a  conical,  flat,  very  prominent  horn;  keel 
veiy  deep  behind  the  shoulders,  conical  and  slightly  inclined  forward,  veined 
along  the  lower  side,  slightly  undulating  and  declining  abruptly  along  half 
its  length,  straight  and  slightly  attenuated  from  thence  to  the  tip,  which  is 
black  and  extends  far  beyond  the  tip  of  the  abdomen;  ridge  mostly  ferrugin- 
ous; sides  slightly  tumid;  wings  colorless;  veins  tawny;  fore-wings  partly 
brown  along  the  hind  borders  and  at  the  tips,  tawny  and  punctured  towards 
the  base.    Length  of  the  body,  4  lines;  of  the  wings,  10  lines. 

New  York. 

Var.  /3.  Head  and  fore  part  of  the  fore-chest  pale  yellow  tinged  with  green ; 
keel  ta\\Tiy,  partly  green;  legs  tinged  with  green;  oviduct  pitchy,  curved. 

St.  John's  Bluff,  E.  Florida. 

Distribution.  Van  Duzee  reports  this  species  from  New  York, 
Pennsylvania  and  Florida.  It  has  also  been  taken  in  Johnson 
county,  Kansas. 

Hosts.    Mrs.  Slosson  and  Van  Duzee  report  it  from  sycamore. 


76  The  University  Science  Bulletin. 

Telamona  tristis  Fitch. 

Telamona  tristis  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  51,  1851. 

Telamona  coryli  Fitch,  Honiop.  N.  Y.  St.  Cab.,  p.  51,  1851. 

Telamona  spreta  Coding,  Bui.  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  417,  1894. 

Funkhouser  gives  the  following  technical  description: 

Near  T.  ampelopsidis  in  appearance,  but  smaller  and  lighter  and  differing  in 
coloration;  crest  high  and  square,  higher  before  than  behind;  tegmina  hyaline 
tipped  with  brown;  pronotum  yellow  mottled  with  red-brown. 

Head  subquadrate,  yellowish,  faintly  longitudinally  striate,  finely  punctate, 
closely  pubescent,  faintly  mottled  with  brown;  eyes  prominent,  brown;  ocelli 
pearly,  nearer  to  each  other  than  to  the  eyes;  clypeus  pubescent,  tip  slightly 
extending  below  inferior  margin  of  face. 

Bronotum  densely  punctate,  not  pubescent,  ground  color  light  yellow,  a 
broad  transverse  reddish-brown  fascia  nearly  covering  metopidium,  a  second 
on  front  of  crest,  and  a  third  extending  down  posterior  third  of  crest  and  reach- 
ing lateral  margin  of  pronotum ;  humeral  angles  produced,  triangular,  flattened, 
blunt,  tips  dark;  dorsal  crest  nearly  square,  truncate  at  tip,  posterior  margin 
pale;  posterior  process  long,  sharp,  not  quite  reaching  tips  of  tegmina. 

Tegmina  smoky  hyaline,  bases  opaque  and  punctate,  tips  brown.  Under- 
surface  of  thorax  flavous;  abdomen  brown.    Legs  ferruginous. 

Length,  8.5  mm.;  width,  5  mm. 

Distribution.  Reported  from  Ontario,  New  York,  New  Jersey, 
Pennsylvania  and  Illinois.  There  are  specimens  in  the  Snow  collec- 
tion from  Kansas  City,  Mo.,  so  it  doubtless  occurs  in  Kansas. 

Hosts.    Funkhouser  reports  this  species  from  hazelnut  and  oak. 

Telamona  ampelopsidis  (Harris). 

Membracis  cissi  (Harris  MS),  List  Ins.  Mass.,  in  Hitchcock,  Geol.  Mass.,  p.  584,  1833. 
Membracis  ampelopsidis  Harris,  Rept.  Ins.  Mass.,  p.  181,  1841. 
Thelia  cyrtops  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  310,  1846. 
Telamona  ampelopsidis  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  51,  1851. 
Telamona  monticola  Uhler,  Stand.  Nat.  Hist.,  ii,  p.  225,  fig.  302,  1884. 

Funkhouser  gives  the  following  technical  description: 

Fine,  large,  well-marked  species;  crest  high,  erect,  front  margin  nearly  per- 
pendicular, hind  margin  sloping;  ground  color  grayish  with  brown  transverse 
fascia  across  metopidium,  deep  brown  area  at  frontal  base,  brown  fascia  extend- 
ing from  posterior  tip  of  crest  to  lateral  margin  of  pronotum;  tegmina  hyaline, 
with  brown  tips. 

Head  yellowish,  faintly  marked  with  brown  below,  sculptured,  finely  punc- 
tate, sparingly  pubescent;  eyes  prominent,  grayish  brown;  ocelli  large,  yellow- 
ish, nearer  to  each  other  than  to  the  eyes;  clypeus  smooth,  pubescent,  tip  tri- 
angular. 

Pronotum  finely  punctate,  very  sparingly  pubescent;  metopidium  yellow  at 
frontal  margin,  black  spot  above  each  eye,  median  carina  prominent,  black; 
humeral  angles  prominent,  blunt,  extending  beyond  the  eyes  as  far  as  the 
length  of  the  eyes;  dorsal  crest  higher  before  than  behind,  margin  somewhat 
flattened;  posterior  process  long,  strong,  heavy,  extending  almost  to  tips  of 
tegmina. 


Lawson:    Membracid.e  of  Kansas.  77 

Teginina  hyaline,  lightly  punctate  at  base  and  along  costal  margins,  tips 
brown.    Undersurface  of  body  generally  uniform  gray-brown. 

Male  smaller  and  darker  than  female,  often  without  characteristic  markings. 

Length:  female,  10  mm.;  male,  8-9  mm.  Width:  female,  6  mm.;  male, 
5  mm. 

Distribution.  This  species  has  been  reported  from  Massachusetts, 
New  York,  New  Jersey,  Maryland,  North  Carolina,  Illinois,  Kansas 
and  Colorado.  It  has  been  taken  in  Kansas  in  Riley,  Leavenworth 
and  Douglas  counties. 

Hosts.  Occurs  only  on  Psedera  quinquefolia,  our  common  Virigina 
creeper. 

Genus  Telamonanthe  Baker. 

The  members  of  this  genus  greatly  resemble  those  of  the  preced- 
ing genus  in  general  appearance,  but  they  are  smaller  insects  as  a 
rule  and  are  all  characterized  by  having  the  basal  costal  cell  of  the 
tegmina  completely  punctate. 

One  species  of  the  genus  has  been  taken  in  Kansas  and  another  at 
Kansas  City,  Mo.,  so  that  both  doubtless  occur  in  the  state. 

These  species  may  be  separated  by  the  following  key: 

KEY   TO   SPECIES. 

A.    Crest  nearly  quadrate,  humeral  angles  very  long.  rileyi. 

A  A.   Crest  rounded,  humeral  angles  short.  modesta. 

Telamonanthe  rileyi  (Coding). 

Telamona  rileyi  Coding,  Ent.  News,  iii,  p.  108,  1892. 

Telamona  coqnilletti  Coding,  Bui.  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  420,  1894. 

Telamotmnlhe  rileyi  Baker,  Can.  Ent.,  xxxix,  p.  115,  1907. 

The  original  description  follows: 

Similar  in  size  and  form  to  coquilletti  Coding;  the  markings  are  less  promi- 
nent, lateral  horns  much  less  produced. 

Male.  Greenish  yellow,  marked  with  ferruginous  lines,  punctured.  Head 
greenish  j'ellow,  lightly  punctured.  Prothorax  yellowish  green,  mottled  with 
ferruginous,  longitudinal,  elevated  lines;  dorsal  carina  percurrent,  a  deeply 
impressed  dot  on  each  side  of  its  base;  dorsal  crest  somewhat  elevated,  much 
compressed,  strongly  compressed  anteriorly  at  base  and  posteriorly  behind 
middle,  the  highest  point  of  crest  at  beginning  of  postesior  third,  from  which 
point  it  gradually  slopes  anteriorly  in  a  gentle  curve  continuous  with  anterior 
third  of  prothorax,  posteriorly  sloping  for  a  short  distance,  then  forming  an 
obtuse  angle;  at  the  base  another  obtuse  angle  is  formed,  from  which  the 
median  carina  curves  gently  to  the  apex;  lateral  angles  a  little  prominent; 
tegmina  with  basal  half  coriaceous,  apical  half  subcoriaceous,  a  brown  spot  at 
apex.  Chest  below  is  dark  yellow,  coxae  piceous.  Legs  yellow  and  hairy; 
abdomen  yellow.    Length,  6  mm. 

Habitat:    Mario  county,  Cal. 


78  The  University  Science  Bulletin. 

Distribution.  The  type  specimen  came  from  California.  There 
are  specimens  in  the  Snow  collection  from  Arizona  and  from  Lincoln 
county,  Kansas. 

Hosts.    Unknown. 

Telamonanthe  modesta  (Goding). 

Telamoim  modesta  Goding,  Bui.   III.   St.   Lab.   Nat.   Hist.,   iii,  p.   420,   1894. 

The  original  description  follows: 

Head  triangular,  hairy;  ocelli  nearer  to  each  other  than  to  the  eyes. 
Prothorax  broad,  convex  in  front,  gradually  elevated  back  of  lateral  angles  in 
a  very  high,  much  compressed,  crest,  the  upper  and  anterior  edges  continuously 
curved  to  base  of  prothorax;  posterior  superior  angle  rectangular,  posterior 
edge  straight,  inclined  forward  somewhat;  posterior  process  long,  depressed, 
acuminate,  gradually  attenuated  to  apex;  sordid  greenish  yellow  covered  with 
black  punctures,  hairy,  two  black  impressed  dots  over  each  eye,  one  above  the 
other;  base  of  posterior  process  and  posterior  edge  of  crest  more  or  less  free 
from  black  punctures.  Tegmina  with  basal  half  of  corium  punctured,  sub- 
transparent.  Legs  triquetrous,  tibiae  punctured  with  black,  covered  with 
spines.    Abdomen  and  chest  greenish-yellow. 

Length.  8  mm.;  breadth  4  mm.;  altitude,  5  mm. 

Habitat :     Galesburg,  111. 

Distribution.  Types  from  Illinois.  There  is  a  specimen  in  the 
Snow  collection  from  Kansas  City,  Mo.,  so  the  species  undoubtedly 
occurs  in  Kansas. 

Hosts.    Unknown. 

Genus  Archasia  Stal. 

The  members  of  this  genus  are  at  once  recognized  by  the  broad, 
leaflike  and  compressed  expansion  of  the  pronotum. 

Van  Duzee  separates  the  two  species  occurring  in  Kansas  by  the 
following  key: 

*  KEY   TO   SPECIES. 

A.   Dorsal  edge  distinctly  brown  or  fuscous,  the  contour  obviously  concave 
before  the  apex.  belfragei. 

AA.   Dorsal  edge  concolorous  or  faintly  dotted  with  brown,  contour  scarcely 
if  at  all  concave  before  the  apex.  galeata. 

Archasia  belfragei  Stal. 

Archasia  belfragei  Stal,  Of.  Vet.   Akad.  Forh.,   xxvi,  p.   250,   1869. 
Archasia  canadensis  Piovancher,  Pet.  Faune  Ent.  Can.,  iii,   p.   230,   1889. 

Funkhouser  gives  the  following  technical  description: 

Green,  fading  to  yellowish  in  cabinet  specimens;  pronotum  high,  strongly 
foliaceous,  dorsal  margin  brown;  tegmina  about  half  concealed  by  pronotum; 
posterior  process  not  reaching  apices  of  tegmina. 

Head  nearly  twice  as  wide  as  long,  smooth,  sparingly'  pubescent;  base  high 
and  sinuate;  eyes  very  prominent,  shining  dark  brown;  ocelh  pearly,  promi- 
nent, nearer  to  each  other  than  to  the  eyes. 


Lawson:    Membracid.e  of  Kansas.  79 

Pronotuni  cioscly  bvit  weakly  piinctatp,  not  pubescent;  humeral  angles 
small,  triangular;  dorsal  crest  very  high,  flattened,  foliaceous,  almost  vertical 
above  head,  slightly  concave  above  head,  posterior  margin  gradually  hollowed 
out  before  apex  of  posterior  process,  entire  dorsal  margin  flattened  and  uni- 
formly brown. 

Tegmina  smoky  iiyaline,  bases  and  costal  margins  punctate,  tips  strongly 
marked  with  brown.  Understirface  of  body  yellow-brown;  abdomen  brown 
Legs  dull  yellow-brown;  tibiae  pubescent. 

Length,  9  mm.;  width,  4.5  mm.;  height  of  pronotum,  5  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario, 
Massachusetts,  New  York,  New  Jersey,  North  Carolina,  Illinois  and 
Michigan.  There  is  a  specimen  in  the  Snow  collection  from  Kansas 
City,  Mo.,  so  it  undoubtedly  occurs  in  Kansas  also. 

Hosts.    Funkhouser  gives  locust  and  oak  as  hosts. 

Archasia  galeata  (Fabricius.) 

Mefnbracis  galeata  Fabricius,  Syst.  Rhyng.,  p.  9,  1803. 

Thelia  yaleata  Fairniaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  309,  1846. 

Smilia  auriciUata  Emmons,  Nat.  Hist.  N.  Y.  Ins.,  p.  153,  pi.  3,  fig.  12,  1854. 

The  following  describes  this  species: 

Green,  fading  to  yellowish  in  cabinet  specimens;  pronotum  very  high  and 
strongh'  foliaceous,  dorsal  margin  concolorous  or  spotted  with  brown;  tegmina 
about  half  concealed  by  pronotimi ;  posterior  process  not  reaching  apices  of 
tegmina. 

Head  shghtly  longer  than  in  preceding  species,  smooth,  sparingly  pubescent; 
base  high  and  sinuate;  eyes  very  prominent,  shining  dark  brown;  ocelli  pearly, 
prominent,  nearer  to  each  other  than  to  the  eyes. 

Pronotum  closely  but  distinctly  punctate,  sparsely  pubescent;  humeral  an- 
gles smaller  and  more  rounded  than  in  preceding  species  and  the  dorsal  crest 
higher,  not  concave  above  the  head,  with  posterior  margin  not  at  all  concava 
before  apex  of  posterior  process,  the  entire  dorsal  margin  strongly  flattened. 

Tegmina  smoky  hyaline,  bases  and  costal  margins  punctate,  tips  slightly 
darker.    Entire  undereurface  and  legs  yellowish-brown,  the  tibiae  pubescent. 

Length,  9-11  mm.;  width,  4.5-5  mm.;  height  of  pronotum,  6  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York,  New  Jersey,  Pennsylvania,  North  Carolina,  Georgia,  Florida, 
Illinois,  Iowa,  Colorado,  Utah  and  Texas.  Specimens  are  at  hand 
from  Douglas,  Riley,  Miami  and  Cowley  counties,  Kansas. 

Hosts.    Coding  mentions  Eupatorium,  Verbena  hastata  and  oak. 

Genus  Smilia  Germar. 

The  members  of  this  genus  are  of  the  same  general  shape  as  those 
of  the  preceding,  because  of  the  foliaceous  dorsum,  but  the  terminal 
cell  of  the  hind  wing  is  not  sessile  and  truncate,  but  petiolate  and 
triangular. 

A  single  species  occurs  in  Kansas. 


80  The  University  Science  Bulletin. 

Smilia  camelus  (Fabricius). 

(PI.   Vll,  figs.    1,   2.) 

Membracis  camelus  Fabricius,   Syst.    Rhyng.,   p.    10,    1803. 

Smilia  vittata  Amyot  &  Sendlle,  Heniip.,  p.   539,   1843. 

Thelia  camelus  Walker,  List  Homop.,  ii,  p.   562,  1851. 

Thelia  vittata  Walker,  List  Homop.,  iv,  p.   1143,    1852. 

Smilia  betulae  Coding,  Can.  Ent.,  xxv,  p.   196,  1893. 

Antianthe  compressa  Buckton,   Monog.   Memb.,  p.   191,  pi.    41,   fig.   6,   1903. 

.  Funkhouser  gives  the  following  technical  description : 

Pronotimi  high  and  foliace'ous,  extending  forward  over  the  head;  brown 
with  broad  diagonal  stripe  of  green  or.  yellowish  followed  by  a  parallel  trans- 
lucent band  and  a  white  spot;  males  much  smaller  and  darker  than  females. 

Head  triangular,  sculptured,  yellow  with  scattered  brown  punctures  and 
hairs;  eyes  brown;  ocelli  pearly,  margins  raised,  nearer  to  each  other  than 
to  the  eyes;  clypeus  continuing  inferior  line  of  face,  apex  slightly  produced. 

Pronotum  coarsely  punctiu-ed,  punctures  farther  apart  in  pale  parts;  wide 
green  band  extending  from  anterior  dorsal  angle  of  crest  to  lateral  margin 
of  pronotum,  this  band  fading  to  yellowish  in  dried  insects;  wide  translucent 
band  from  behind  middle  of  dorsum  to  lateral  base  of  crest;  white  spot  at 
posterior  base  of  crest;  humeral  angles  hardly  produced,  short,  rounded; 
posterior  process  short,  pointed,  not  reaching  tips  of  tegmina. 

Tegmina  hyaline,  bases  punctate  with  brown,  apices  brown.  Undersur- 
face  of  body  brownish  yellow.    Legs  flavous. 

Length:  female,  9  mm.;  male,  7-8  mm.  Width:  female,  3  mm.;  male, 
2.5-3  mm. 

Internal  male  genitalia.  Styles  with  anterior  portions  broadened 
and  flattened  as  in  Telamona,  the  sparsely  spined  posterior  portions 
of  nearly  same  width  to  the  laterally  curved  and  doubly  toothed 
apices;  connective  large,  broad,  much  as  in  Telamona;  oedagus, 
viewed  laterally,  large,  U-shaped,  anterior  arm  lightly  chitinized 
and  longer  than  the  more  heavily  chitinized  posterior  arm,  with  a 
distinct  knob  at  base  and  also  about  midway  up  the  anterior  arm, 
the  posterior  arm  with  functional  orifice  on  caudal  aspect  of  apex 
and  covered  with  teeth  on  the  cephalic  aspect. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
Hampshire,  Massachusetts,  New  York,  New  Jersey,  Pennsylvania, 
North  Carolina,  Georgia,  Florida,  Illinois,  Michigan,  Iowa,  Mis- 
souri, and  Texas.  Specimens  have  been  taken  in  Kansas  in  Douglas 
and  Montgomery  counties. 

Hosts.  Funkhouser  gives  locust  and  oak  as  hosts. 

Genus  Cyrtglgbus  Goding. 
The  members  of  this  genus  are  usually  brownish  insects  without 
prominent  humeral  angles,  with  a   compressed  dorsum,  and   fre- 
quently with  a  thin,  semitransparent  spot  below  the  dorsal  ridge. 


Lawson:    Membracid.e  of  Kansas.  81 

Tlie  genus  is  represented  by  at  least  nine  species  in  Kansas,  which 
live  on  various  species  of  oaks. 

Van  Duzee  divides  the  genus  into  four  subgenera  which  he  sepa- 
rates by  the  following  key : 

KEY  TO   SUBGENERA. 

Pronotum  strongly  inflated  posteriorly,  the  crest  forming  an  inflated  cyst  be- 
fore and  behind  the  median  pale  spot.  Xantholobus. 

Pronotum  not  strongly  inflated,  the  crest  however,  usually  more  or  less  dis- 
tinctly compressed  on  the  median  spot 1 

1.  Dorsal  crest  low  and  sinuated  at  the  middle;   form  elongated  and 
much  depressed.                                                                 Evashmeadea. 
Dorsal  crest  not  distinctly  sinuated  above 2 

2.  Crest  highest  anteriorly  over  or  before  the  humeral  angles,  the  dor- 
sum sloping  and  nearly  rectilinear  to  the  apex.  Atymna. 
Crest  regularly  arcuated  above  in  the  more  typical  examples,  or  in 
any  case  with  the  highest  point  at  or  near  the  middle. 

Cyrtolobus. 

Subgenus  Cyrtolobus  Coding. 

KEY  TO  SPECIES. 

A.   Dorsum  without  anterior  notch  or  depression. 

B.   Crest  well  developed.  celsiis. 

BB.   Crest  distinctly  lower.  fenestratus. 

AA.   Dorsum  with  anterior  depression  before  elevation. 

B.   Crest  arising  before  humeral  angles.  fuliginosus. 

BB.   Crest  arising  behind  humeral  angles. 

C.   Large  species,  at  least  9  mm.  in  length.  tuberosus. 

CC.   Small  species,  not  over  7  mm.  in  length. 

D.    Crest  low  or  obsolete. 

E.   Species  distinctly  brownish.  griseus. 

EE.   Species  distinctly  greenish.  cinereus. 

DD.    Crest  well   developed;    pronotum  with   distinct   oblique 
bands.  vau. 

Cyrtolobus  celsus  Van  Duzee. 

Cyrtolobus  celsus  Van  Duzee,  Check  List  Hemip.,  p.  60,  1916  (n.  n.  for  fenestratus 
Van  D.). 

Cyrtolobus  fenestratus  Van  Duzee,  Bui.  Buf.  Soc.  Nat.  Sci.,  ix,  p.  81,  1908. 

The  following  is  the  original  description: 

Pronotum  well  elevated,  highest  at  about  the  middle,  almost  regularly  arcu- 
ated from  base  to  tip,  the  dorsal  line  a  very  little  broken  at  the  posterior  vitta. 
Head  and  the  pronotum  anterior  to  the  oblique  line  pale  yellowish  testaceous, 
punctured  and  varied  with  brown;  from  above  each  eye  a  brown  indefinite 
vitta  curv^es  backward  over  the  humeral  angle;  anterior  oblique  vitta  almost 
perpendicular  at  first,  approaching  the  elongated  median  mark,  then  deflected 
and  again  widened  at  apex  so  as  to  pass  almost  straight  across  the  dorsal 
carina ;  posterior  vitta  transverse,  represented  in  my  Atlanta  specimen  only  by 
a  narrow  spot  on  the  carina;  the  surface  behind  the  anterior  vitta  darker,  in 
one  example  almost  piceous.    Elytra  hyaline,  the  smoky  apex  small  and  palo. 


82  The  University  Science  Bulletin. 

Face  coarsely  punctured;  clypeus  broad,  moderately  produced,  and  strongly 
incurved. 

Length,  6  mm.;  height,  3  mm. 

Distribution.  Van  Duzee  reports  this  species  from  Massachu- 
setts, New  York,  New  Jersey  and  Georgia.  It  has  also  been  taken 
near  Kansas  City,  Mo.,  and  therefore  undoubtedly  occurs  in  Kansas. 

Cyrtolobus  fenestratus  (Fitch). 

Cyrtosia  fenestrata  Fitch,  Hoinop.  N.  Y.  St.   Cab.,   p.   49,  1851. 

Cyrotolobus  fenestratus  Goding,  Can.  Ent.,  .xxv,  p.   172,   1893. 

Cyrtolobus  muticus  Van  Duzee,  Bui.  Buf.  Soc.  Nat.  Sci.,  ix,  p.  83,  1908  (in  part). 

The  following  is  the  original  description: 

Yellow  marbled  with  rufous;  a  pellucid  spot  behind  the  summit  of  the  keel 
and  a  smaller  one  halfway  to  the  apex;  an  oblique  yellow  vitta  below  the  an- 
terior spot,  margins  with  fuscous  or  sanguineous;  tip  of  the  thorax  reaching 
beyond  the  terminal  cells  of  the  elytra.  Male  black,  the  pellucid  spots  almost 
obsolete  and  the  yellow  vitta  replaced  by  a  few  yellow  dots. 

Length,  0.25.    On  oaks. 

Distribution.  Van  Duzee  reports  this  species  from  Massachu- 
setts, New  York,  New  Jersey,  Pennsylvania,  North  Carolina,  Geor- 
gia, Florida,  Mississippi,  Ohio,  Illinois,  Colorado,  Dakota.  Speci- 
mens have  been  taken  at  Kansas  City,  Mo.,  so  it  surely  occurs  in 
Kansas. 

Cyrtolobus  fuliginosus  (Emmons). 

Cyrtosia  fuliginosa  Emmons,  Nat.  Hist.  N.  Y.  Ins.,  p.  154,  pi.   13,  fig.   15,  1854. 
Cyrtolobus  fuliginosus  Goding,  Can.  Ent.,  xxv,  p.   172,  1893. 

Funkhouser  gives  the  following  technical  description: 

Near  C.  ovatus  in  appearance,  but  smaller,  darker,  and  with  lower  crest; 
dark  sordid  brown  with  faint  transverse  bands;  head  projecting  slightly  for- 
ward; posterior  process  just  reaching  tips  of  tegmina;  tegmina  strongly 
marked  with  brown,  apices  lighter. 

Head  somewhat  extended  forward,  yellow,  mottled  with  deep  brown,  deeply 
punctate  with  brown,  not  pubescent,  a  black  spot  at  base  of  head  above  each 
ocellus;  eyes  large,  brown,  lighter  in  color  than  remainder  of  head;  ocelli 
small,  pearly,  about  equidistant  from  each  other  and  from  the  eyes;  clypeus 
convex,  sculptured,  a  brown  line  on  each  side,  tip  continuing  rounded  inferior 
outline  of  face. 

Pronotum  dark  brown,  transverse  fascia  extending  from  anterior  base  of 
crest  to  lateral  margin  of  pronotum,  this  fascia  light  browTi  before  and  very 
dark  brown  behind;  entire  pronotum  deeply  and  densely  punctate;  humeral 
angles  weak,  angular  but  blunt;  dorsal  crest  regularly  arcuate  from  above 
humeral  angles  to  base  of  posterior  process;  posterior  process  heavy,  short, 
blunt,  just  reaching  apices  of  tegmina. 

Tegmina  smoky  brown,  apical  cells  lighter,  apical  margins  fuscous,  bases 
and  costal  margins  roughly  punctate.    Legs  and  undersurface  of  body  flavous. 

Length,  6  mm.;  width,  2.5  mm. 


Lawson:    Mkmbracid.e  of  Kansas.  83 

Distribution.  Hitherto  reported  only  from  New  York.  Specimens 
have  been  taken,  however,  at  Kansas  City,  Mo.,  and  so  it  may  safely 
be  included  in  the  Kansas  fauna. 

Hosts.    Funkhouser  reports  it  from  white  oak. 

Cyrtolobus  tuberosus  (Fairmaire). 

Thelia  tuberosus  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  307,   1846. 
Cyrtolobus  tuberosus  Goding,  Bui.  III.  St.  Lab.  Nat.  Hist.,  iii,  p.  433,  1894. 

Funkhouser  gives  the  following  technical  description: 

Largest  species  of  the  genus;  brown  mottled  with  darker  brown;  dorsal 
compression  strikingl}-  transparent;  dorsal  crest  situated  well  back  on  prono- 
tuni.  posterior  process  very  short;  tegmina  smoky  hyaline  tipped  with  brown. 

Head  triangular,  broader  than  long,  ochraceous  tinged  with  red  and  punc- 
tate with  brown,  not  pubescent;  base  weakly  sinuate;  inferior  margin  of  face 
strongly  sinuate;  eyes  large,  brown;  ocelli  small,  yellowish,  slightly  protrud- 
ing, nearer  to  each  other  than  to  the  eyes;  clj'peus  convex,  brown  line  on  each 
side,  tip  extended  and  hairy. 

Pronotum  deeply  and  closely  punctate,  light  greenish  brown;  crest  dark 
brown  with  pale  compression  at  anterior  base,  in  the  middle,  and  at  posterior 
base;  middle  compression  veiy  large  and  transparent,  posterior  half  of  crest 
dark  brown  with  color  extending  in  a  dark  band  to  margin  of  pronotum; 
metopidium  very  convex,  median  carina  prominent;  humeral  angles  promi- 
nent, rounded;  posterior  process  short,  sharp,  brown,  inferior  lateral  margin 
slightly  sinuate,  not  reaching  tips  of  tegmina. 

Tegmina  brownish  h3-aline,  tips  strongly  marked  with  brown,  bases  punc- 
tate. Undersurface  of  thorax  yellow.  Legs  ferruginous,  hind  trochanters 
marked  with  brown;  tarsi  flavous;  claws  brown. 

Length,  9.5  mm.;  width,  4  mm.  Male  smaller  than  female,  but  similarly 
colored. 

Distribution.  Reported  by  Van  Duzee  from  Massachusetts,  New 
York,  North  Carolina,  Georgia,  Florida,  Mississippi  and  Illinois. 
There  is  a  specimen  in  the  Snow  collection  from  Kansas  City,  Mo., 
so  it  is  sure  to  occur  in  Kansas. 

Hosts.    Funkhouser  reports  it  from  red  oak  and  hickory. 

Cyrtolobus  griseus  Van  Duzee. 

Cyrtolobus  griseus  Van  Duzee,  Bui.  Buf.  Soc.  Nat.  Sci.,  ix,  p.  90,  1908. 

The  following  is  the  original  description : 

This  form,  which  is  very  near  ciner'eus,  is  of  an  almost  uniform  dark  cinerous, 
closely  punctured  with  fuscous.  The  only  conspicuous  marking  is  the  oblique 
blackish  vitta  which  in  cinercus  bounds  the  hind  margin  of  the  anterior  oblique 
vitta.  Before  and  behind  this  brown  band  or  line  the  surface  is  a  little  lighter 
and  there  is  a  suggestion  of  the  median  dorsal  spot  and  transverse  posterior 
vitta.  In  some  specimens  there  is  an  arcuated  longitudinal  brown  vitta  on 
either  side  of  the  metopidium.  Here  the  elytra  are  hyaline  with  the  costal 
base  ferruginous  grey  and  punctured,  and  the  apex  has  a  small  fuscous  cloud. 


84  The  University  Science  Bulletin. 

The  tergum  is  black  with  a  broad  white  band  at  the  base  as  in  the  alHed  spe- 
cies, which,  showing  through  the  closed  elytra,  indicates  the  markings  found 
on  the  elytra  of  cinereus.  Face  obviously  longer  and  more  convex  than  in 
cinereus,  with  the  basal  middle  depressed,  and  the  clypeus  and  lorse  together 
larger,  the  former  much  broader,  more  convex  and  more  decurved  and  rounded 
at  apex. 

Distribution.    The  type  material  was  taken  at  Effingham,  Kansas. 

Cyrtolobus  cinereus  (Emmons). 

Gargara  cinereus  Emmons,  Nat.  Hist.  N.  Y.  Ins.,  p.  156,  pi.  13,  fig.  3,  1854. 

Cyrtosia  ornata  Provancher,  Pet.  Faune  Ent.  Can.,  iii,  p.  240,  1889. 

Cyrtosia  cinerea  Harrington,  Ottawa  Nat.,  vi,  p.  30,  1892. 

Cyrtolobus  cinereum  Coding,  Can.  Ent.,  xxv,  p.  172,   1893. 

Atymna  cinereum  Coding,  Bui.  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  436,  1894. 

Cyrtolobus  cinereus  Van  Duzee,  Bui.  Buf.  Soc.  Nat.  Sci.,  ix,  p.  91,  1908. 

Funkhouser  gives  the  following  technical  description: 

Small  greenish  gray  mottled  with  brown  and  banded  with  green;  pronotum 
low  and  regularly  arcuate;  metopidium  convex;  posterior  process  short  but 
sharp;  tegmina  wrinkled,  hyaline,  apices  brown. 

Head  convex,  pale  grayish  green,  sharply  punctate  with  black,  sparingly 
pubescent;  base  nearly  straight;  eyes  prominent,  brown;  ocelU  large,  reddish, 
prominent,  slightly  farther  from  each  other  than  from  the  eyes  and  situated 
sUghtly  below  an  imaginary  line  extending  through  centers  of  eyes;  clypeus 
flat,  somewhat  trilobed,  a  faint  brown  line  on  each  isde,  extending  below  in- 
ferior margin  of  face. 

Pronotum  green-gray  tinged  with  reddish,  closely  punctate,  not  pubescent; 
dorsal  crest  very  low,  median  spot  on  margin  pale ;  a  transverse  pale  band  bor- 
dered with  brown  extending  from  anterior  base  of  crest  backward  and  down- 
ward to  lateral  margin  of  pronotum,  a  similar  band  extending  from  base  of 
posterior  process  downward  and  forward  to  almost  meet  the  anterior  stripe 
and  form  a  V  with  it ;  posterior  process  short,  not  reaching  tips  of  tegmina. 

Tegmina  wrinkled,  hyaline,  brown  spot  at  base  of  each,  another  in  middle, 
and  a  third  at  tip;  areas  between  hyaline.  Legs  and  undersurface  of  body 
grayish  flavous. 

Length,  5.8  mm.;  width  2.5  mm. 

Distribution.  Reported  by  Van  Duzee  from  Quebec,  New  York 
and  New  Jersey.  A  single  specimen  of  this  species  was  taken  in 
Douglas  county,  Kansas,  and  is  in  the  Snow  collection.  Specimens 
have  also  been  taken  in  Riley  county. 

Cyrtolobus  i^au  (Say). 

(PI.  Vll,  figs.  5,  6.) 

Membracis  van  Say,  Jl.  Acad.  Nat.  Sci.  Phila.,  vi,  p.  299,  1831. 

Thelia  seinifascia  Walker,  List  Homop.,  ii,  p.   561,   1851. 

Smilia  vau  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  48,  1851. 

Thelia  vau  Walker,  List  Homop.,  iv,  p.  1142,  1852. 

Cyrtosia  vau  Provancher,  Pet.  Faune.  Ent.  Can.,  iii,  p.  238,   1889. 

Cyrtosia  jenestrata  Provancher,  Pet.  Favme  Ent.  Can.,  iii,  p.  239,  1889. 

Cyrtolobus  nigra  Coding,  Can.  Ent.,  xxv,  p.  172,  1893. 

Cyrtolobtis  punctifrontis  Coding,  Can.   Ent.,   xxv,  p.   172,   1893- 


Lawson:    Membracid^  of  Kansas.  85 

Cyrtolobus  tricincta  Coding,  Can.  Ent.,  xxv,  p.  172,  1893. 

Cyrtolobus  van  Coding,  Can.  Ent.,  xxv,  p.  172,  1893. 

Thelia  fasciata  Buckton,  Monog.  Membr.,  p.  189,  1893. 

Argante  semifasciata  Buckton,  Monog.  Membr.,  p.  189,  pi.  40,  fig.  9,  pi.  41,  fig.  1,  1903. 

Cyrtolobus  variM  Smith,  Cat.  Ins.  N.  J.,  edn.  3,  p.  92,  1910. 

Funkhouser  gives  the  following  technical  description: 

Small  robust  species,  with  low  pronotum  and  prominent  markings;  varies 
greatlj'  in  color  and  somewhat  in  size;  females  larger  and  lighter  than  males, 
but  with  constant  markings;  transverse  pronotal  band  prominent,  pale  bor- 
dered with  deep  brown;  dorsal  compression  deep  and  translucent;  posterior 
process  short,  blunt,  not  reaching  tips  of  tegmina;  tegmina  hyaline,  with  bases 
and  tips  slightly  brown. 

Head  small,  subtriangular,  pale  yellow  punctured  with  brown;  base  feebly 
sinuate;  inferior  margin  of  face  sinuate,  c^'peus  extending  slightly  below  line; 
eyes  large,  gray-brown;  ocelli  small,  yellowish,  somewhat  nearer  to  each  other 
than  to  the  eyes;  clypeus  hairy. 

Pronotum  closely  and  roughly  punctate;  median  compressed  spot  round, 
transparent;  dorsal  crest  low,  arising  above  humeral  angles  and  gradually  ex- 
tending with  only  a  faint  sinus  before  posterior  process;  posterior  process  short, 
blunt,  tectiform,  reaching  to  bases  of  apical  cells  of  tegmina. 

Tegmina  hyaline,  veins  prominent,  bases  and  apices  smoky  hyaline.  Legs 
and  undersurface  of  body  uniform  flavous. 

Length,  5.5-6.5  mm.;  width,  2.4-2.6  mm. 

Internal  male  genitalia.  Styles  rather  long  and  slender,  curved 
strongly  laterad  apically,  and  bearing  several  spiny  tubercles  on  the 
lateral  margins  just  cephalad  of  the  terminal  tooth;  connective  tri- 
angular; cedagus,  viewed  laterally,  large,  U-shaped,  anterior  arm 
slightly  chitinized  and  with  a  large  prominence  about  half  way  on 
its  cephalic  aspect,  posterior  arm  heavily  chitinized,  apex  with  file- 
like teeth  on  the  side  opposite  the  functional  orifice. 

Distribution.  Reported  by  Van  Duzee  from  Ontario,  Maine,  New 
Hampshire,  Massachusetts,  New  York,  New  Jersey,  Pennsylvania, 
District  of  Columbia,  North  Carolina,  Georgia,  Florida,  Illinois, 
Kansas,  Missouri,  Arkansas,  Texas,  New  Mexico  and  Colorado. 
There  are  specimens  in  the  Snow  collection  from  Douglas,  Riley,  and 
Pottawatomie  counties,  Kansas. 

Hosts.  Funkhouser  gives  white,  chestnut,  red  and  scarlet  oak 
as  hosts. 

Subgenus  Atymna  Stal. 

In  this  subgenus  the  pronotum  is  highest  at  the  anterior  end. 
It  is  represented  in  Kansas  by  a  single  species. 

Cyrtolobus  querci  (Fitch). 

(PI.   VII,  figs.   3,   4.) 

Smilia  querci  Fitch,  Homop.  N.  Y.  St.  Cab.,  p.  49,  1851. 

Thelia  querci  Walker,  List  Homop.,  iv,  p.   1143,  1852. 

Gargara  querci  Emmons,  Nat.  Hist.  N.  Y.  Ins.,  p.   156,  pi.  13,  fig.  8,  1854. 


86  The  University  Science  Bulletin. 

Atymna  querci  Van  Duzee,  Psyche,  v,  p.  390,  1890. 

Cyrtolohus  {Atymna)  querci  Van  Duzee,  Bui.  Buf.   Soc.  Nat.   Sci.,  v,  p.   188,   1894. 

Funkhouser  gives  the  following  technical  description: 

Females  large  and  green,  males  smaller  and  brown  with  a  broken  yellow 
median  dorsal  stripe;  body  long  and  narrow;  crest  highest  above  humeral 
angles  and  gradually  sloping  to  posterior  apex  without  a  sinus. 

Female.  Head  projecting  slightly  forward,  pale  yellow,  sculptured,  irregu- 
larly punctate,  not  pubescent;  eyes  very  prominent,  reddish;  ocelli  not  promi- 
nent, yellow;   clypeus  extending  below  inferior  margin  of  face. 

Pronotum  uniform  green,  roughly  punctate,  not  pubescent,  dorsal  line 
faintly  marked  with  brown;  posterior  process  short,  acute,  not  reaching  tips 
of  tegmina. 

Tegmina  entirely  hyaline,  bases  and  costal  margins  faintly  punctate;  hind 
wings  iridescent.    Legs  and  imdersurface  of  body  green. 

Length,  7  mm.;  width,  2.5  mm. 

Male.  Head  sordid  yellow,  sculptured,  sparingly  punctate;  eyes  prominent, 
brown;  ocelli  pearly;  clypeus  marked  with  brown  at  base. 

Pronotum  chocolate  brown  with  bright  yellow  stripe  on  median  dorsal  line 
and  yellow  band  before  apex. 

Tegmina  smoky  hyaline  with  brown  cloud  at  apices.  Undersurface  of 
thorax  brownish;  abdomen  xevy  dark  brown,  nearly  black.  Legs  flavous;  tarsi 
ferruginous;  claws  fuscous. 

Length,  6  mm.;  width,  2  mm. 

Internal  male  genitalia.  Styles  small,  anterior  portion  quite  short, 
posterior  portions  larger,  bearing  spiny  protuberances  apically ;  con- 
nective rather  large,  triangular;  oedagus,  viewed  laterally,  very  large 
compared  with  the  styles,  anterior  arm  long  and  slender,  posterior 
arm  stout  and  with  many  filelike  teeth  on  the  side  opposite  the 
functional  orifice. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  Con- 
necticut, New  York,  Pennsylvania,  North  Carolina,  Georgia,  Illi- 
nois, Michigan,  Iowa  and  Missouri.  Specimens  having  been  taken 
at  Kansas  City,  Mo.,  it  is  sure  to  occur  in  Kansas. 

Subgenus  Xantholobus  Van  Duzee. 

In  this  subgenus  the  pronotum  is  distinctly  inflated  posteriorly. 
It  is  represented  in  Kanas  by  a  single  species. 

Cyrtolohus  muticus  (Fabricius). 

Membracis  mutica  Fabricius,  Genera  Ins.,  p.  297,  1776. 

Cicada  mutica  Gmelin  in  Linnaeus,  Syst.   Nat.,   edn.   13,   i,  pt.   4,  p.   2093,   1778. 

Centrotus  mutica  Fabricius,   Syst.  Rhyng.,   p.    21,   1803. 

Membracis  trilineatus  Say,  Long's  2nd.  Exped.,  p.   300,  1824;    Compl.   Writ.,  i,  p.   200. 

Cyrtosia  mutica  Stal,   Heniip.  Fabr.,  ii,  p.   25,   1869. 

Cyrtosia  trilineata  Provancher,  Pet.  Faune  Ent.  Can.,  iii,  p.  239,   1889. 

Cyrtolobus  muticus  Coding,  Bui.   111.   St.   Lab.  Nat.   Hist.,  iii,  p.   431,   1894. 


Lawson:    Membracid^  of  Kansas.  87 

Funkhouscr  gives  the  following  technical  description: 

Yellowish  tinged  with  red;  transverse  band  of  pronotuni  often  absent; 
pronotum  long;  head  slightly  projecting  forward;  eyes  tinged  with  reddish; 
posterior  process  reaching  tijxs  of  tegniina ;  teginina  entirely  hyaline  or 
faintlj'  clouded  with  yellow. 

Head  slightly  protruding  forward,  j-ellow  with  red  punctures,  sculptured, 
not  pubescent;  base  irregularly  sinuate;  eyes  gray  marked  with  red;  ocelli 
small,  translucent,  somewhat  nearer  to  each  other  than  to  the  eyes;  clypeus 
swollen,  convex,  continuing  inferior  outline  of  face,  tip  slightly  extended, 
hairy;   antenme  prominent. 

Pronotum  yellowish  with  irregular  reddish  areas,  deeply  and  roughly 
punctate,  not  pubescent;  transverse  band  when  present  pale  with  reddish 
borders;  humeral  angles  weak,  blunt;  dorsal  crest  elliptical,  very  slight  sinus 
before  base  of  posterior  process,  compressions  not  deep;  posterior  process 
heavj-,  blunt,  just  reaching  tips  of  tegmina. 

Tegmina  hyaline  or  clouded  with  reddish  yellow,  tips  pale,  veins  in  some 
cases  yellowish,  bases  and  costal  areas  irregularly  punctate.  Legs  and  under- 
surface  of  thorax  flavous;  abdomen  sordid  yellow. 

Length,  6  mm.;  width,  2.8  mm. 

Distribution.  Reported  hy  Van  Duzee  from  Quebec,  Rhode 
Island,  New  York,  Pennsylvania  and  Illinois.  In  the  Snow  col- 
lection there  are  specimens  from  Kansas  City,  Mo.,  and  some  taken 
in  Douglas  county,  Kansas.  The  writer  took  a  specimen  at  St. 
Paul,  ]\Iinn. 

Genus  Ophiderma  Fairmaire. 

Tlie  members  of  this  genus  have  a  compressed  and  rounded  dor- 
sum which  entirely  lacks  a  crest. 

It  is  represented  in  Kansas  by  three  species,  which  may  be  sep- 
arated by  the  following  key: 

KEY   TO   SPECIES. 

A.    Color,  browTi  or  mottled. 

B.    Species  smaller,  mottlings  more  distinct.  salamandra. 

BB.   Species  larger,  mottling  dull.  flaviguttula. 

AA.   Color,  green  or  yellowish  green.  flava. 

Ophiderma  salamandra  Fairmaire. 

(PI.  VI,  figs.   \  2.) 
Ophiderma  salamandra  Fairmaire,  Ann.  Soc.  Ent.  Fr.,  ser.  2,  iv,  p.  493,  1846. 

Funkhouser  gives  the  following  technical  description: 
Large  brown  species;  dorsum  rounded  and  very  pubescent  with  short,  black, 
bristly  hairs;  posterior  process  short,  suddenly  acute,  not  reaching  apices  of 
tegmina;  tegmina  hyaline,  bases  and  costal  areas  strongly  punctate,  tips 
clouded  with  fuscous,  veins  very  prominent ;  under  part  of  body  dark ;  males 
smaller  and  darker  than  females. 


88  The  University  Science  Bulletin. 

Head  broader  than  long,  yellow,  feebly  punctate,  very  hairy;  base  slightly, 
uniformly  curved;  eyes  large,  bro^vn;  ocelli  prominent,  red,  nearer  to  each 
other  than  to  the  eyes;  inferior  margin  of  face  sinuate;  clypeus  yellow  with 
two  vertical  stripes  of  red;  base  hairy. 

Pronotum  coarsely  punctate,  densely  pubescent,  brown  mottled  with  green; 
dorsum  rounded,  slightly  depressed  behind  middle,  lateral  margin  curved 
downward  at  middle;  posterior  process  short,  suddenly  acute,  not  reaching 
tips  of  tegmina. 

Tegmina  smoky  hyline,  veins  very  prominent,  nearly  all  of  basal  half  below 
pionotum  strongly  punctate,  tips  clouded  with  fuscous;  hind  wings  iridescent. 
Undersurface  of  head  and  thorax  fuscous;  abdomen  flavous.  Femora  and 
tibiae  strongly  marked  with  dark  brown. 

Length,  7.6  mm.;  width,  3.2  mm. 

Internal  male  genitalia.  Styles  rather  short  and  broad,  especially 
basally,  posterior  portion  more  slender,  apices  curved  strongly 
laterad  and  ending  in  a  small  but  distinct  terminal  hook  on  the 
lateral  margin,  in  front  of  which  is  a  distinct  prominence  which 
tapers  gradually  to  the  hook,  the  apical  fourth  of  the  styles  bearing 
a  few  spines;  connective  almost  pentagonal,  with  a  pair  of  distinct 
knobs  at  the  basal  angles;  oedagus,  viewed  laterally,  U-shaped, 
anterior  process  with  a  large  anterior  prominence  near  the  middle, 
posterior  arm  curved,  the  apex  bearing  the  functional  orifice  caudally 
and  many  filelike  and  distinct  teeth  on  its  cephalic  aspect. 

Distribution.  Reported  by  Van  Duzee  from  Ontario,  New  Hamp- 
shire, Massachusetts,  New  York,  New  Jersey,  Pennsylvania,  District 
of  Columbia,  Virginia,  Georgia,  Florida,  Illinois  and  Michigan. 
There  are  specimens  in  the  Snow  collection  also  from  Wisconsin, 
Arizona  and  Kansas  City,  Mo.  It  has  also  been  taken  in  Potta- 
watomie county,  Kansas. 

Hosts.    Oaks. 

Ophiderma  jiaviguttula  Coding. 

Ophiderma  jiaviguttula  Coding,  Bui.  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  438,  1894. 
Ophiderma  flavoguttata  Slosson,   Ent.    News,  xvii,   p.   326,   1906. 

The  original  description  follows: 

Female.  Head  triangular,  yellowish;  eyes  prominent,  dark  brown;  ocelli 
equidistant  from  each  other  and  from  the  eyes,  red;  convex,  densely  pubes- 
cent. Prothorax  with  very  slight  median  carina,  densely  pubescent,  an 
irregular  yellow  patch  starting  at  lateral  border  and  extending  upwards  and 
forwards,  midway  between  base  and  apex;  an  irregular  band  at  base,  con- 
colorous  with  head,  extending  along  sides  in  a  greenish  gray  line;  otherwise 
dirty  brown,  lightly  punctvired;  apex  of  posterior  process  not  reaching  apex 
of  tegmina.  Tegmina'  subcoriaceous  at  base,  lightly  punctured,  basal  half 
and  apex  brown.    Below  yellow,  feet  and  legs  brown. 

Length,  6.2  mm. 


Lawson:    Membracid^  of  Kansas.  89 

Distribution.  Reported  by  Van  Duzee  from  New  Hampshire, 
Pennsylvania  and  Illinois.  There  arc  specimens  in  the  Snow  col- 
lection from  Kansas  City,  Mo.,  so  it  undoubtedly  occurs  in  Kansas. 

Hosts.  Probably  oak. 

Ophiderma  fiava  Goding. 

Ophiderma  fiava  Goding,  Insect  Life,  v,  p.  93,  1892  (nomen  nudum). 
Ophiderma  fiava  Goding,  Bui.  111.  St.  Lab.  Nat.  Hist.,  iii,  p.  439,  1894. 

Funkhouser  gives  the  following  technical  description: 

Large  greenish  yellow  species,  fading  to  sordid  yellow  in  cabinet  specimens; 
body  robust  and  long;  posterior  process  not  reaching  apices  of  tegmina; 
tegmina  h3'aline,  brown  at  base  and  fuscous-clouded  at  tips. 

Head  much  broader  than  long,  green,  weakly  and  sparingly  punctate, 
smooth,  shining,  sparingly  pubescent;  eyes  large,  red;  ocelli  prominent,  red- 
dish, about  equidistant  from  each  other  and  from  the  eyes;  clypeus  smooth, 
nearh^  black,  base  regularh'  rounded,  tip  extending  below  inferior  margin  of 
face. 

Pronotum  uniform  green,  in  some  cases  tinged  with  reddish,  closely  and 
densel}^  punctate,  finely  pubescent;  dorsum  rounded,  depressed  behind  middle, 
median  carina  percurrent;  posterior  process  heavy,  tectiform,  acute,  not  ex- 
tending to  tips  of  tegmina. 

Tegmina  hyaline,  bases  reddish  and  punctate,  tips  clouded  with  fuscous, 
veins  heavy  and  inclined  to  be  punctate  along  margin.  Legs  and  undersur- 
face  of  body  entirely  flavous. 

Length,  7-8  mm.;  width,  3-4  mm. 

*  Distribution.     Reported  by  Van  Duzee  from  Quebec,  Massa- 
chusetts, New  York,  Pennsylvania,  North  Carolina,  Illinois,  Michi- 
gan.   There  are  specimens  in  the  Snow  collection  from  Columbia, 
Mo.,  and  from  Chautauqua  county,  Kansas. 
Hosts.   Unknown. 

Genus  Vanduzea  Goding. 
The  members  of  this  genus  are  distinguished  by  the  transverse 
and  basally  truncate  terminal  cell  of  the  elytra. 
A  single  member  of  the  genus  is  known  to  occur  in  Kansas. 

Vanduzea  triguttata  (Burmeister). 

(PI.  VI,  figs.  7,  8.) 

Entylia  triguttata  Burmeister,  Silb.  Rev.  Ent.,  iv,  p,  183,  1836. 

Vanduzea  vestita  Goding,  Insect  Life,  v,  p.  93,  1892. 

Cyrtolobus  annexus  (Uhl.  MS)  Townsend,  Can.  Ent.,  xxiv,  p.   196,  1892. 

The  following  is  Coding's  description : 

Head  broad,  black,  perpendicular,  triangular,  a  narrow  dusky  brown  mark 
on  upper  edge  just  below  origin  of  carina.  Eyes  prominent;  ocelli  equidistant 
from  each  other  and  the  eyes.  Front  of  prothorax  blackish  brown,  fading 
posteriorly  to  a  reddish  brown  in  a  triangular  form,  the  apex  of  which  reaches 


90  The  University  Science  Bulletin. 

three-fourths  of  the  distance  to  apex  of  posterior  process ;  lateral  angles  slightly 
produced;  sides  of  prothorax  from  just  behind  lateral  angles  to  apex  black, 
interrupted  by  a  light  yellow,  or  whitish,  trapezoidal  spot  on  each  side  just 
behind  middle  of  inferior  border;  just  before  the  apex  a  white  band  across 
posterior  part  of  process.  Tegmina  clear,  with  dark  brown  veins,  or  brown 
with  a  lighter  band  across  middle.  Legs  and  feet  brown  or  .black. 
Length,  4.7  mm. 

Internal  male  genitalia.  Styles  broad  and  flat  basally,  apical  half 
slender,  apices  curved  strongly  dorsad  and  ending  rather  bluntly, 
bearing  a  few  spines ;  connective  nearly  quadrangular,  the  base  con- 
cave; oedagus,  viewed  laterally,  U-shaped,  anterior  arm  enlarged 
toward  the  base,  posterior  arm  of  nearly  same  width  throughout, 
the  apex  bearing  caudally  the  functional  orifice  and  cephalad  many 
filelike  teeth. 

Distribution.  Reported  by  Van  Duzee  from  District  of  Columbia, 
Florida,  Colorado,  New  Mexico  and  Arizona.  The  writer  has  taken 
it  at  St.  Paul,  Minn.  It  has  been  taken  in  the  following  Kansas 
counties:  Morton,  Clark,  Stevens,  Logan,  Haskell,  Seward,  Riley, 
Hamilton,  Pottawatomie,  Douglas  and  Miami. 

Hosts.  Common  on  black  locust.  Also  taken  by  the  writer  on 
A7norpha. 

Genus  Entylia  Germar. 

The  members  of  this  genus  are  characterized  by  their  high  com- 
pressed dorsum,  which  bears  a  deep  median  notch. 

A  single  species  of  the  genus  is  known  to  occur  in  Kansas. 

Entylia  concisa  Walker. 

(PI.  VI,  figs.  5,  6.) 

Entylia  concisa  Walker,  List  Homop.,  ii,  p.   547,  1851. 

Entylia  decisa  Walker,  List  Homop.,  ii,  p.  548,  1851. 

Entylia  concava  Piovancher,  Pet.  Faune  Ent.  Can.,  iii,  p.   233,  1889. 

The  following  is  the  original  description: 

Ferruginea;  prothoracis  carina  alte  bicristata,  utrinque  albo  interrupte  et 
oblique  fasciata ;  pedes  flavi ;  alse  limpidfe ;  alse  anticse  basi  et  ad  costam  fulvae. 

Ferruginous;  head  and  thorax  roughly  punctured;  head  transverse,  almost 
semicircular,  narrower  than  the  fore-chest,  slightly  impressed  with  an  indis- 
tinct middle  suture  which  extends  to  the  face,  the  hind  border  of  the  latter  is 
angular  and  occupies  nearly  half  the  length  of  the  head;  shoulders  very  ob- 
tusely angular,  not  prominent:  fore-chest  forming  two  lofty  compressed  keel- 
shaped  crests  which  inchne  towards  each  other  and  inclose  three-fourths  of  a 
circle;  the  first  rises  between  the  shoulders  and  is  truncated  at  the  tip;  the 
second  Is  lower  and  above  the  keel;  the  latter  is  rather  deep  and  extends  far 
beyond  the  tip  of  the  abdomen,  whose  sides  it  embraces;  the  irregular  ridges 
on  the  sides  of  the  crest  communicate  with  the  ridges  of  the  keel,  and  the  lat- 
ter has  an  obUque  white  interrupted  band  on  each  side  behind  the  second 


Lawson:    Membracid.e  of  Kansas.  91 

crest;  breast  jiitchy;  abdomen  black;  a  stripe  on  each  side  beneath  and  the 
tip  yellow ;  legs  yellow ;  wings  colorless ;  fore-wings  tawny  at  the  base  and  along 
more  than  half  the  length  of  the  fore  border;  veins  pitchy,  tawny  where  the 
wings  are  so,  pale  yellow  near  the  tip  of  the  fore  border. 

Length  of  the  abdomen,  2%  hnes;  of  the  wings,  4%  lines. 

Var.  /)' .    Breast  and  abdomen  tawny;  white  bands  of  the  keel  hardly  visible. 

Var.  y.    Breast  and  abdomen  black;  tip  of  the  latter  tawn3\ 

St.  John's  Bluff,  E.  Florida. 

Internal  male  genitalia.  Styles  small,  anterior  portion  narrow, 
posterior  part  stouter,  the  apices  curved  strongly  laterad  and  ending 
in  a  stout  hook;  connective  large,  triangular,  apex  abtuse,  and  longi- 
tudinally divided;  oedagus,  viewed  laterally,  U-shaped,  anterior  arm 
with  a  distinct  protuberance,  posterior  arm  stouter  and  ending  in  a 
large,  serrated  point. 

Distribution.  Van  Duzee  reports  this  species  from  District  of 
Columbia,  North  Carolina,  Georgia  and  Florida.  It  has  been  taken 
in  Douglas,  Pottawatomie  and  Wilson  counties,  Kansas. 

Remarks.  Funkhouser  feels  that  E.  sinuata  is  the  very  small 
southern  form,  E.  bactriana  the  northeastern,  and  that  E.  concisa  is 
our  Kansas  species.  In  the  present  confused  status  of  the  members 
of  this  genus  it  is  perhaps  best  to  accept  his  conclusions  and  to 
change  our  Kansas  species  to  E.  concisa  instead  of  calling  it  E.  sin- 
uata, as  did  Miss  Branch. 

Hosts.  Miss  Branch  reports  this  species  from  Melilotus  alba, 
Cnicus  altissinius,  Phleum  alpinum,  Helianthus  annuus,  Medicago 
sativa,  and  Ambrosia  sp.  The  writer  has  taken  it  very  commonly  in 
all  its  stages  from  Ambrosia  triflda,  Xanthium  sp.,  and  Helianthus 
tuberosus. 

Genus  Publilia  Stal. 

The  members  of  this  genus  are  closely  related  to  those  of  the  pre- 
ceding genus,  but  have  a  much  lower  crest  and  a  much  weaker  me- 
dian notch. 

All  of  the  members  of  the  genus  found  in  the  United  States  occur 
in  Kansas.  These  may  be  separated  by  the  following  key  given  by 
Van  Duzee. 

KEY  TO  SPECIES. 

A.   Dorsum  straight  or  feebly  arcuated,  scarcely  if  at  all  sinuated;  form  more 
slender.  mudesta. 

AA.   Dorsum  more  elevated,  obviously  sinuated. 

B.    Sides  of  the  pronotum  with  longitudinal  rugae  which  become  more 
or  less  reticulated  along  the  dorsum.  concava. 

BB.   Rugae  of  the  pronotum  strong,  irregularly  reticulated  over  its  whole 
surface.  reticulata. 


92  The  University  Science  Bulletin.  "^  ^^ 

Publilia  modesta  Uhler. 

(PI.  VI,  fig.  3,  4.) 

Publilia  modesta  Uhler,  Bui.   U.   S.  Geol.   Geog.   Surv.,  i,  p.    .344,   1876. 
Publilia  bicinctura  Coding,  Ent.  News,  iii,  p.  200,  1892. 

The  original  description  follows: 

General  form  of  P.  concaua  Say,  but  more  decidedly  vertical  in  front,  and 
with  the  dorsal  outline  scarcely  depressed  before  the  middle;  apex  of  the  pro- 
notum  more  slender  and  acute.  Color  pale  yellow;  the  head  and  fore  part  of 
pronotum  clouded  with  pale  brown;  a  spot  above  the  humeri,  a  broad  obhque 
band  behind  the  middle,  and  a  broad  cloud  on  the  apex  grayish  brown;  the 
dorsal  edge  irregularly  spotted  with  dark  brown,  and  the  oblique  band  sur- 
mounted by  a  large  brown  spot.  The  surface  closely  beset  with  series  of  coarse 
sunken  jjunctures;  the  longitudinal  and  reticulated  surface  lines  obsolete. 
Humeral  margin  of  the  sinus  waved.  Under  side  piceo-testaceous;  the  front, 
clypeus,  pectu.s  and  venter,  excepting  the  edges  of  the  segments,  black-piceous. 
Legs  dull  yellow,  closely  pubescent,  clouded,  and  spotted  with  brown. 

Length,  4  to  4%  mm.;  breadth  of  pronotum,  2  mm. 

Colorado  (C.  Thomas) ;  also  discovered  in  Utah,  Dakota,  Arizona,  New 
Me.xico,  Cahfornia.  , 

In  two  specimens  examined,  the  commonly  raised  lines  on  the  surface  of  the 
pronotum  were  obliterated. 

Distribution.  Besides  the  above  localities  specimens  have  been 
taken  in  Kansas  in  Gove  and  Trego  counties. 

Hosts.  Gillette  and  Baker  report  this  species  on  Solidago,  alfalfa, 
Helianthus,  Iva,  and  Artemesia.  Goding  gives  Glycyrrhiza  lepidota 
and  mesquite  as  hosts. 

Publilia  concava  (Say). 

Membracis  concava  Say,  Long's  2nd  Exped.,  ii,  p.  301,  1824;   Compl.   Writ.,  i,  p.  200. 

Entylia  concava  Germar,  Silb.  Rev.  Ent.,  iii,  p.   249,  1835. 

PublUia  concava  Stal,  .\nalecta  Hem.,  p.   388,   1866. 

Ceresa  concava  Rathvon,  in  Mombert's  Hist.  Lane.  Co.,  Pa.,  p.   5.51,  1869. 

Publilia  grisea  Buckton,  Monog.  Membr.,  p.   184,  pi.  39,  fig.  5,  1903. 

Publilia  vittata  Buckton,  Monog.  Membr.,  p.  185,  pi.  39,  fig.  6,  1903. 

Funkhouser  gives  the  following  technical  description: 

Varies  greatly  in  color  and  somewhat  in  shape,  particularly  in  form  of  dor- 
sal sinuation;  color  varies  from  gray  to  black;  dorsum  convex,  tectiform, 
faintly  ribbed,  dorsal  sinus  shallow;  pronotum  irregularly  ridged,  deeply  punc- 
tate; tegmina  largely  co\ered  by  pronotum,  basal  half  of  each  costal  area 
strongly  punctate. 

Head  slightly  projecting,  strongly  punctate  with  black;  base  nearly  straight; 
inferior  margin  I'ounded;  ej'es  not  prominent;  ocelli  prominent,  usually  red- 
dish; clypeus  rounded,  very  wide  at  tip. 

Pronotum  deeply,  densely  and  coarsely  punctate,  lateral  areas  marked  with 
high,  distinct,  irregular,  longitudinal  ridges;  dorsal  margin  sinuate  just  be- 
hind humeral  angles,  sinuation  usually  very  shallow;  posterior  lobe  gradually 
elliptical  to  posterior  apex;  posterior  process  heavy,  high,  tectiform,  blunt,  ex- 
tending just  beyond. tips  of  tegmina. 


Lawson:    Membracid.e  of  Kansas.  93 

Tegmina  almost  entirely  concealed  by  pronotum ;  exposed  costal  margins 
opaque  and  punctate  for  basal  half,  ajncal  areas  hyaline,  tips  fuscous.  Under- 
surface  of  body  and  femora  usually  very  dark,  generally  black.    Legs  flavous. 

Length,  5  mm.;  width,  2.5  mm. 

Internal  male  genitalia.  Styles  long  and  slender,  the  extreme  an- 
terior portion  bent  strongly  laterad,  widest  opposite  connective,  the 
posterior  part  with  distinct  apical  hook;  connective  widest  at  the 
concave  base,  the  apex  truncate;  oedagus,  viewed  laterally,  with 
strong  anterior  arm  which  bears  two  prominent  rounded  promi- 
nences, the  posterior  arm  more  slender,  the  apical  portion  bearing  a 
few  indefinite  teeth  on  caudal  margin  and  the  apex  characteristically 
retrorsely  hooked. 

Distribution.  Van  Duzee  reports  this  species  from  Ontario,  New 
York,  New  Jersey,  North  Carolina,  Ohio,  Illinois,  Iowa,  Missouri, 
Kansas,  Arkansas,  and  Utah.  It  is  also  known  to  occur  in  Mexico. 
Specimens  have  been  taken  in  Kansas  in  Douglas,  Pottawatomie, 
Riley,  and  Gove  counties. 

Hosts.  Funkhouser  gives  golden  rod  and  skunk  cabbage  as  hosts. 
Goding  mentions  Canadian  thistle.  The  writer  has  taken  it  in  all 
stages  on  Ambrosia  trifida. 

Publilia  reticulata  Van  Duzee. 

Pxiblilia  reticulata  Van  Duzee,  Bui.  Buf.  Soc.  Nat.   Sci.,  ix,  p.   106,  1908. 

The  following  is  the  original  description : 

Closely  allied  to  concava  but  with  the  surface  of  the  pronotum  reticulated 
with  strong  anastomosing  rugae  in  place  of  the  four  or  five  simple  longitudinal 
carinse  found  in  that  species.  These  rugae  give  the  surface  a  strongly  corru- 
gated or  areolated  appearance.  Surface  between  the  rugae  deeply  punctured. 
Metopidiimi  more  vertical  than  in  concava,  the  percurrent  carina  more  ele- 
vated and  the  dorsal  sinus  somewhat  deeper.  Apical  margin  of  the  head  dis- 
tinctly sinuated  next  the  eye,  then  very  obtusely  arcuated  about  the  apex. 
Color  blackish  or  fuscous,  speckled  or  blotched  more  or  less  with  testaceous  on 
the  head  and  front  of  the  pronotum  as  far  as  the  dorsal  sinus;  and  with  a  tri- 
angular whitish  testaceous  spot  on  the  apical  fourth  of  the  lateral  margin, 
which  may  be  prolonged  somewhat  along  the  margin  anteriorly  and  indis- 
tinctly across  the  disk  as  an  incomplete  subapical  vitta.  Venter,  tibiae  and 
tarsi  pallid,  the  disk  of  the  ventral  segments  more  or  less  black. 

Length,  4  mm. 

Distribution.  Van  Duzee  reports  this  species  from  New  Jersey, 
Pennsylvania,  North  Carolina  and  Missouri.  There  is  a  specimen 
in  the  Snow  collection  from  Kansas  City,  Mo.,  and  it  has  been  taken 
in  Riley  county. 

Hosts.    Unknown. 


94  The  University  Science  Bulletin. 


PLATE  II. 

1.  Styles  and  coqgctive  of  Ceresa  bubalus. 

2.  ffidagus  of  Ceresa  bubalus. 

3.  Lateral  aspect  of  Ceresa  taurina. 

4.  Genitalia  of  Ceresa  taurina. 

5.  Styles  and  connective  of  Ceresa  bubalus. 

6.  ffidagus  of  Ceresa  bubalus. 

7.  Genitalia  of  Ceresa  bubalus. 

8.  Lateral  aspect  of  Ceresa  bubalus. 


Lawson:    Membracid^  of  Kansas. 


95 


PLATE  11. 


96  The  University  Science  Bulletin. 


PLATE  III. 

1.  Genitalia  of  Ceresa  dicer os. 

2.  ffidagus  of  Ceresa  diceros. 

3.  Styles  and  connective  of  Ceresa  palnieri. 

4.  ffidagus  of  Ceresa  palmeri. 

5.  Styles  and  connective  of  Stictocephala  jestina. 

6.  Lateral  aspect  of  Stictocephala  jestina. 

7.  ffidagus  of  Ceresa  borealis. 

8.  Genitalia  of  Ceresa  borealis. 


Lawson:    Membracid.e  of  Kansas. 


97 


PLATE  IIT. 


98  The  University  Science  Bulletin. 


PLATE  IV. 

1.  Genitalia  of  Stictocephala  inermis. 

2.  CEdagus  of  Stictocephala  inermis. 

3.  Genitalia  of  Stictocephala  lutea. 

4.  ffidagus  of  Stictocephala  lutea. 

5.  Genitalia  of  Stictocephala  inermis. 

6.  Qildagus  of  Stictocephala  inermis. 

7.  Lateral  aspect  of  Stictocephala  inermis. 

8.  Styles  and  connective  of  Stictocephala  inermis. 


Lawson:    Membracid.e  of  Kansas. 


99 


PLATE  IV. 


100  The  University  Science  Bulletin. 


PLATE  V. 

1.  Lateral  aspect  of  genitalia  of  Thelia  bimaculata. 

2.  Dorsal  aspect  of  genitalia  of  Thelia  bimaculata. 

3.  Dorsal  aspect  of  genitalia  of  Micrutalis  calva. 

4.  Lateral  aspect  of  genitalia  of  Micrutalis  calva. 

5.  Dorsal  aspect  of  genitalia  of  Acutalis  tartarea. 

6.  Lateral  aspect  of  genitalia  of  Acutalis  tartarea. 

7.  Lateral  aspect  of  Telamona  pyramidata. 

8.  Styles  and  connective  of  Telamona  pyramidata. 


Lawson:    Membracid.e  of  Kansas. 


101 


PLATE  V. 


102  The  University  Science  Bulletin. 


PLATE  VI. 

1.  Lateral  aspect  of  genitalia  of  Ophiderma  salamandra. 

2.  Dorsal  aspect  of  genitalia  of  Ophiderma  salamandra. 

3.  Lateral  aspect  of  genitalia  of  Publilia  modesta. 

4.  Dorsal  aspect  of  genitalia  of  Publilia  modesta. 

5.  Dorsal  aspect  of  genitalia  of  Entylia  concisa. 

6.  Lateral  aspect  of  genitalia  of  Entylia  ccmcisa. 

7.  Dorsal  aspect  of  genitalia  of  Vanduzea  triguttata. 

8.  Lateral  aspect  of  genitalia  of  Vanduzea  triguttata. 

9.  Lateral  aspect  of  genitalia  of  Campylenchia  latipes. 

10.  Ventral  aspect  of  genitalia  of  Campylenchia  latipes. 

11.  Dorsal  aspect  of  genitalia  of  Enchenopa  binotata. 

12.  Lateral  aspect  of  genitalia  of  Enchenopa  binotata. 


Lawson:    Membracid.e  of  Kansas. 


103 


PLATE  VI. 


104  The  University  Science  Bulletin. 


PLATE  VII. 

1.  Lateral  aspect  of  genitalia  of  Smilia  camelus. 

2.  Dorsal  aspect  of  genitalia  of  Smilia  camelus. 

3.  Dorsal  aspect  of  genitalia  of  Cyrtolobus  querci. 

4.  Lateral  aspect  of  genitalia  of  Cyrtolobus  querci. 

5.  Dorsal  aspect  of  genitalia  of  Cyrtolobus  vau. 

6.  Lateral  aspect  of  genitalia  of  Cyrtolobus  vau. 

7.  Dorsal  aspect  of  genitalia  of  Telamona  viridia. 

8.  Lateral  aspect  of  genitalia  of  Telamona  viridia. 


Lawson:    Membracid.e  of  Kansas. 


105 


PLATE  VII. 


INDEX. 

V.\C,  K 

ac'uiniiuitus,  tJki.s.suiiutus 67 

Acutalis ()2 

tartarea 62 

albescens,  Ceresa. 52 

Archasia 78 

l)elfragei 78 

galeata 79 

Key  to  species 78 

Atynina 85 

belfragei,  Archasia 78 

bimaculata,  Thelia 66 

binotata,  Enchenopa 49 

l)()realis,  Ceresa 54 

brevicornis,  Ceresa 55 

bubalus,  Ceresa 57 

calva,  JMicrutalis 63 

camelus,  Smilia 80 

Campylenchia 47 

latipes 47 

carvff,  Microcentrvis .  .  .' 45 

Carynota 64 

mera 64 

celsus,  Cyrtoloyjus 81 

Centrotinae 45 

Ceresa 51 

albescens.  . 52 

liorealis 54 

l)revicornis 55 

bubalus 57 

diceros 51 

Key  to  species 51 

palmeri 53 

taurina 56 

cinereus,  Cyrt<)Iol)Us 84 

collina,  Telamona 75 

concava,  Pulililia 92 

concisa,  Entylia 90 

cristata,  Heliria 68 

Cyrt  Globus 80 

celsus 81 

cinereus 84 

fenestratus 82 

f  uliginosus 82 

griseus 83 

Key  to  subgenera 81 

muticus 86 

querr'i    85 

(107) 


108  The  University  Science  Bulletin. 


PAGE 


Subgenus 81 

Key  to  species 81 

tuberosus 83 

vau 84 

decorata.  Telamona 72 

diceros,  Ceresa " 51 

Distribution 32 

Economic  importance 43 

Enchenopa 48 

binotata 49 

Entylia 90 

eoncisa 90 

extrema,  Telamona 74 

fenestratus,  Cyrtolobus 82 

festina,  Stictocephala 61 

flava,  Ophiderma 89 

flaviguttula,  Ophiderma 88 

f uliginosus,  Cyrtolobus 82 

_galeata,  Archasia 79 

Genitalia,  male 38 

Glossonotus 67 

acuminatus 67 

griseus,  Cyrtolobus 83 

Heliria 68 

cristata 68 

Key  to  species 68 

scalaris ._^ 68 

inermis,  Stictocephala 59 

latipes,  Campylenchia 47 

Life  history 41 

List  of  species '  .  .  .  .  44 

lugubris,  Telamona 72 

lutea,  Stictocephala ; 60 

Membracins 47 

Key  to  genera 47 

mera,  Carynota 64 

Microcentrus 45 

carya3 45 

Micrutalis 63 

calva 63 

modesta,  Pul)lilia 92 

modesta,  Telamonanthe 78 

muticus,  Cyrtolobus 86 

obsoleta,  Telamona 71 

Ophiderma 87 

flava 89 

flaviguttula 88 

Key  to  species 87 

salamandra 87 


Lawson:    Membracid^  of  Kansas.  109 

PAGE 

palineri.  Ceresa 53 

Phylogeny 41 

Puialilia 91 

concava 92 

Key  to  species 91 

modesta ' 92 

reticulata 93 

pyramidata,  Tclainona 70 

querci,  Cyrtolobus 85 

querci,  Telamona 73 

reticulata,  Publilia 93 

rileyi,  Telamonanthe 77 

salamandra,  Ophiderma 87 

scalaris,  Heliria 68 

Smilia 79 

cainelus 80 

Smiliinae 50 

Key  to  genera 50 

Stictocephala 59 

festina 61 

inermis 59 

Key  to  species 58 

lutea ■ 60 

Structural  characteristics 36 

Subfamilies,  Key  to .• 45 

tartarea,  Acutalis 62 

taurina,  Ceresa 56 

Telamona 69 

collina 75 

decorata 72 

extrema 74 

Key  to  species 69 

lugubris 72 

obsoleta 71 

pyramidata 70 

querci .  73 

unicolor 74 

viridia 71 

Telamonanthe 77 

Key  to  species 77 

modesta 78 

rileyi 77 

Thelia 65 

biniaculata 66 

Key  to  species 65 

uhleri 65 

triguttata,  Vanduzea 89 

tuberosus,  Cyrtolobus 83 

uhleri,  Thelia 65 


110  The  University  Science  Billetin. 

PAOE 

unicolor,  Telamona 74 

Vancluzea 89 

triguttata 89 

vail,  Cyrtolobus , .  84 

viridia,  Telamona 71 

Xantholobus 86 


THE 

KANSAS  UNIVEKSITY 

Science  Bulletin 


Vol.  XIV,  No.  4— October,  1922. 

(Whole  Series,  Vol.  XXIX,  No.  4.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

The  Genus  Acinopterus  (Homopter.\) P.  B.  Lawson. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  4. 


The  Genus  Acinopterus  (Homoptera,  Cicadellidae) , 

BY  PAUL  B.  LAWSON, 
Professor  of  Entomology,   University  of  Kansas. 


INTRODUCTION. 

THE  genus  Acinoptenis  was  erected  by  Van  Duzee  in  1892  to 
accommodate  the  species  Acinoptenis  acuminatus.  Later,  in 
1895,  Baker  described  a  single  specimen  as  Phlepsius  inornatus,  a 
species  which  Van  Duzee  in  his  catalogue  lists  as  Acinopterus  acu- 
minatus var.  inornatus.  Finally,  in  1903,  Ball  described  three  varie- 
ties of  the  typical  species  of  the  genus,  the  first  of  which,  variety 
variegatus,  Van  Duzee  made  a  synonym  of  Baker's  variety  inorna- 
tus. Thus  up  to  the  time  that  the  writer  undertook  this  study,  one 
species  and  three  varieties  were  recognized  as  comprising  the  mem- 
bership of  this  genus.  The  results  of  the  study  show  that  we  now 
have  at  least  eight  species  and  one  variety.  One  or  two  other  species 
are  probably  represented  in  the  material  at  hand,  but  because  of  the 
scarcity  of  specimens  it  was  not  thought  best  to  describe  them  as  new 
species. 

The  writer  is  indebted  to  the  following  workers  for  very  kindly 
loaning  him  the  material  studied :  Mr.  E.  L.  Dickerson,  Mr.  George 
G.  Ainslie,  Mr.  F.  H.  Lathrop,  Prof.  J.  G.  Sanders,  Dr.  Dwight 
M.  De  Long,  Mr.  C.  E.  Olson,  Mr.  Edmund  H.  Gibson,  Prof.  H.  0. 
Osborn,  and  Dr.  E.  D.  Ball.  The  work  done  is  based  for  the  most 
part  on  the  many  specimens  from  the  collection  of  Doctor  Ball,  who 
not  only  loaned  him  all  his  own  material,  but  also  the  types  in  the 
collection  of  the  Iowa  State  Agricultural  College,  and  in  addition 
was  a  great  help  in  making  suggestions  and  giving  information  as 
to  distribution  and  host  plants.  To  Mr.  E.  H.  Gibson  and  Mr.  W. 
L.  McAfee  the  writer  owes  the  privilege  of  examining  the  collection 

(113) 


114  The  University  Science  Bulletin. 

of  the  National  Museum,  including  the  type  of  Baker's  inornatus. 
Through  the  kindness  of  Dr.  J.  Chester  Bradley,  the  material  at 
Cornell  University  was  studied,  and  Prof.  C.  P.  Gillette  kindly  per- 
mitted the  examination  of  the  collection  of  the  Colorado  Agricul- 
tural College. 

DISTRIBUTION. 

As  far  as  our  present  knowledge  goes,  the  members  of  this  genus 
have  hitherto  been  taken  only  in  the  United  States  and  Mexico,  a 
single  specimen  at  least  having  been  collected  as  far  south  as  Yuca- 
tan. In  our  country  it  is  for  the  most  part  southern  and  western  in 
its  distribution.  This,  along  with  its  occurrence  in  Mexico,  would 
indicate  its  probable  presence  throughout  Central  America  and  per- 
haps in  the  northern  portion  of  South  America.  For  this  last  state- 
ment, however,  there  is  no  certain  data.  The  genus  is,  therefore, 
both  Nearctic  and  Neotropical. 

The  following  states  have  yielded  specimens  of  this  genus:  New 
Jersey,  Maryland,  District  of  Columbia,  North  Carolina,  South 
Carolina,  Virginia,  West  Virginia,  Florida,  Alabama,  Mississippi, 
Tennessee,  Arkansas,  Missouri,  Kansas,  Oklahoma,  Colorado,  Utah, 
Washington,  California,  Arizona,  Texas  and  Georgia. 

HOST  PLANTS. 

Very  little  information  is  available  as  to  the  specific  plants  on 
which  the  members  of  this  genus  feed.  Most  of  the  specimens 
collected  were  taken  while  sweeping  grasses  or  weeds.  De  Long 
reports  Acinopterus  acuminatus  as  abundant  on  grasses  in  Tennes- 
see. Ainslie  took  specimens  from  blue  grass.  Doctor  Ball  believes 
that  A  viridis  var.  variegatus  feeds  on  wild  geranium.  A  single 
specimen  of  A.  angulatus  was  swept  from  Sphoeralcea  angustifolia. 
A  label  on  a  specimen  of  A.  acuminatus  from  Oklahoma  states  that 
it  was  taken  from  Amphiachyris,  while  another  specimen  from 
Rosser,  Tex.,  was  taken  from  Tetraneuris.  Evidently  most  of  the 
members  of  the  genus  are  either  grass  feeders  or  else  have  as  their 
hosts  various  weeds  found  in  grassland.  Much  more  careful  collect- 
ing will  need  to  be  done  before  we  can  be  certain  of  the  specific 

hosts. 

DESCRIPTION  OF  THE  GENUS. 

The  following  is  Van  Duzee's  description  of  the  genus: 

General  appearance  of  Allygus,  but  with  the  elytra  strongly  narrowed 
posteriorly,  and  the  tip  acute. 

Head  narrower  than  the  pronotum,  rounded,  or  somewhat  produced  before, 


Lawson:    Genus  Acinopterus.  115 

with  the  apex  subacute,  hind  edge  broadly  concave.  Vertex  rather  short, 
sloping,  convex  or  more  or  less  impressed  behind  the  apex,  surface  punctured, 
the  anterior  submargin  obscurely  transversely  rugose,  passage  to  the  front 
roimded.  Front  rather  broad,  at  the  base  slightly  encroaching  upon  the  apex 
of  the  vertex,  suddenh^  narrowed  at  tip.  Clypeus  much  widened  apically. 
Lorae  large.  Cheeks  wide.  Pronotum  broad  and  rather  short,  anterior  edge 
broadly  arcuate,  posterior  nearly  straight;  sides  long,  oblique,  carinated; 
lateral  angles  prominent,  latero-posterior  rounded.  Scutcllum  rather  small. 
Elytra  narrow,  lanceolate  at  apex,  the  tip  acute,  appendix  wanting;  sutural 
edge  straight  to  the  extreme  tip;  costal  and  apical  margins  continuous;  apical 
areoles  five,  inner  small,  oblique,  second  largest,  reaching  the  extreme  tip; 
third  and  fourth  small;  fifth,  or  stigmatal,  long  and  usually  crossed  by  one  or 
two  transverse  veinlets;  first  and  second  sectors  united  by  but  one  transverse 
nervure,  but  there  are  usually  three  or  four  connecting  the  outer  claval  nervure 
with  the  claval  suture;  all  the  nervures  strong;  costa  feebly  convex.  Super- 
numerary cell  of  the  wings  present.    Otherwise  as  in  Athysanus  and  Allygus. 

The  above  description  needs  modification  to  enable  it  to  include 
the  several  species  in  which  the  tegmina,  while  visibly  narrowed 
apically,  do  not  end  in  an  acute  tip,  the  sutural  margin  in  these 
cases  not  continuing  on  straight  to  the  extreme  apex,  but  meeting 
the  costal  margin  to  form  a  rather  broadly  rounded  tip.  The  writer, 
therefore,  proposes  the  following  generic  description: 

Head  usually  narrower  than  pronotum,  rarely  as  wide.  Vertex  rounded  or 
distincth'  angulate,  usually  impressed  behind  the  apex,  sloping,  broadly  round- 
ing with  the  front.  Front  broad  basally,  much  narrowed  apically.  Clypeus 
widened  apically.  Lorse  large,  nearly  reaching  margin  of  the  wide  gense  ven- 
trally.  Pronotum  wide,  over  twice  as  wide  as  long,  anterior  and  posterior 
margins  nearly  parallel,  latter  usually  more  nearly  straight;  lateral  and 
humeral  margins  distinct;  disc  transversely  wrinkled.  Scutellum  finely  granu- 
lar and  with  distinct  transverse  impressed  line.  Tegmina  moderately  long,  apex 
alwaj's  narrowed,  sometimes  to  an  acute  tip.  Venation  distinct.  Of  the  five 
apical  cells  M4  is  the  largest,  but  it,  along  with  Cui  and  Ro,  is  frequently 
divided  by  cross-veins,  R2  being  so  regularly  divided  that  probably  both  Ri 
and  R2  are  present.  Cell  2d  M  is  not  divided  by  a  cross-vein.  Veins  1st  A  and 
2d  A  almost  always  connected  by  one  or  more  cross-veins.  Valve  of  male 
always  hidden  under  last  ventral  segment;  plates  usually  long  but  never 
equaling  pygofers.  Female,  last  ventral  segment  large,  always  bearing  a  median 
notch. 


116  The  University  Science  Bulletin. 

SYSTEMATIC  TREATMENT  OF  THE  SPECIES. 

KEY  TO   SPECIES. 

A.    Apex  of  tegmina  strongly  acute;  costal  margin  straight  clear  to  tip. 
B.   Brownish  or  greyish  species. 

C.   Darker  and  larger  species. 

D.  Male  plates  long  and  narrow,  parallel-margined,  apices 
rounded;  last  ventral  segment  of  female  without  lateral 
angles.  acuminatus  Van  D. 

DD.  Male  plates  broad  and  shorter,  apices  distinctly  diver- 
gent; last  ventral  segment  of  female  with  distinct  lateral 
angles.  angulatus  n.  sp. 

CC.    Lighter  and  more  slender  species.  inornatus  (Bak.). 

BB.    Greenish  species.  productus  n.  sp. 

AA.   Apex  of  tegmina  rounded;  costal  margin  not  running  straight  clear  to  tip. 
B.    Species  brownish  or  greyish,  not  distinctly  green. 

C.   Species  large;    last  ventral  segment  of   female  strongly  pro- 
duced medially.  brunneus  Ball. 
CC.    Species    smaller;    last    ventral    segment    of    female    normally 
produced. 

D.    Species  with  elytra  distinctly  variegated. 

viridis  var.  variegatus  Ball. 

DD.   Species  with  elytra  not  variegated  pallidus  n.  sp. 
BB.    Greenish  species. 

C.   Species  larger;  plates  of  male  tapering  and  distinctly  divergent 
apically.  viridis  Ball. 

CC.   Species   smaller;    plates    of    male    parallel-margined   and   but 
slightly  divergent  apically.  obtutus  n.  sp. 

Acinopterus  acuminatus  Van  D. 

(PI.  VIII,  fig.   1;    pi.  IX,  fig.   1;    pi.  X,  fig.    1;    pi.  XI,  fig.   1;    pi.   XII,  figs.   1,   2.) 
Acinopterus  acuminatus  Van  D.,   Psyche,   vi,   p.    308,    1892. 
Acinopterus  acuminatus  G.  &  B.,  Heniip.  Colo.,  p.  94,  1895. 

Acinoptervs  acuminatus  Van  D.,  Bui.  Buf.  Soc.  Nat.  Sci.,  viii.  No.  5,  p.  69,  1907;  ix,  p. 
225,  1909. 

Acinopterus  acuminatus  Osb.,  Ohio  Nat.,  ix,  p.   466,   1909. 
Acinopterus  acuminatus  Smith,  Cat.  Ins.  N.   J.,  edn.   3,  p.   105,   1910. 
Acinopterus  acuminatus  Barb.,  Bui.  Am.  Mus.  Nat.   Hist.,  xxxiii,  p.  534,   1914. 
Acinopterus  acuminatus  Van  D.,  Trans.   San  Diego  Soc.  Nat.   Hist.,  ii,  p.   54,   1914. 
Acinopterus  acuminatus  Mete,  Jl.  Elisha  Mitchell   Sci.   Soc,  xx.xi,  p.   23,   1915. 
Acinopterus  acuminatus  De   L.,   Tenn.   St.    Bd.   Ent.,   Bui.    17,   p.    89,   1916. 
Acinopterus  acujninatus  Van  D.,  Cat.  Hemip.  N.  A.,  p.  675,  1917. 
Acinopterus  acuminatus  Lathr.,   S.   C.  Agr.  Exp.   Sta.,  Bui.   199,  p.    102,   1919. 
Acinopterus  acuminatus  Laws.,  Kans.   Univ.   Sci.   Bui.,  xii,  p.   207,  1920. 

The  following  is  the  original  description  of  this  species: 

Fulvous  brown  tinged  with  dull  green  or  yellowish,  elytral  nervures  pale, 
brown-margined.    Length,  5-6%  mm. 

Head  pale.  Front  with  about  eight  brown  arcs,  more  or  less  distinct. 
Pronotum  feebly  calloused  on  the  anterior  margin,  with  a  few  shallow  im- 
pressions arranged  parallel  to  the  edge,  more  obvious  in  the  males;  lateral 
margin  as  long  as  the  latero-posterior,  acutely  carinated;  disc  posteriorly  ob- 
scurely wrinkled.  Basal  angles  of  the  scutellum  with  a  brownish  triangular 
spot  more  or  less  apparent.  Pectoral  pieces  usually  more  or  less  invaded  with 
blackish,    sometimes    pale    and    immaculate.      Legs    pale,    or    suffused    with 


Law  son:    Genus  Acinopterus.  117 

sanguineous.  Abdomen  pale,  frequently  black  above,  excepting  the  broad 
lateral  margins;  infuscated  on  the  basal  and  apical  segments  of  the  venter  in 
the  females;  slightly  suffused  with  a  pale  median  line  in  the  males.  Elytra 
pale  fulvous,  frequently  whitish  hyaline  on  the  disc  of  the  costal  and  some  of 
the  discal  areoles  of  the  corium,  and  on  the  inner  margin  of  the  clavus,  the 
extreme  apex  clouded  with  smoky  or  even  blackish;  nervures  pale,  edged 
with  brownish,  the  marginal  with  a  fuscous  interruption  at  tip;  claval  suture 
brown.    Wings  smok}^  iridescent,  nervures  fuscous. 

Genitalia.  Male:  Valve  wanting.  Plates  long  and  narrow,  a  little  longer 
than  the  last  ventral  segment,  about  one-third  wider  at  base  than  at 
their  obtusely  lanceolate,  divergent  tips.  Pygofers  twice  the  length  of  the 
plates,  narrowed  and  obtusely  pointed  at  apex,  armed  beyond  the  plates  with 
numerous  stout  spines.  Female:  Last  ventral  segment  rather  long,  hind  edge 
with  a  shallow  median  notch,  either  side  of  which  is  a  broadly  roimded  lobe, 
retreating  at  the  outer  angles.  Pygofers  rather  broad,  their  subacute  apex 
moderately  exceeded  by  the  oviduct. 

Described  from  5  males,  3  females.  Maryland,  September  29  and  August  4, 
on  pines  (Uhler) ;  North  Carolina  (Osborn) ;  New  Jersey  (Uhler) ;  mountains 
of  northwest  Colorado  (Gillette);  California  (Coquillett). 

In  a  male  from  California  the  lower  surface  of  the  femora  is  black.  A 
female  from  North  Carolina  has  the  disc  of  the  elytra  white-pruinose,  and  all 
the  specimens  exhibit  considerable  variation  in  the  extent  of  the  black 
markings. 

The  writer  gives  the  following  description : 

A  large,  rather  robust,  brownish  species,  ranging  from  a  greyish-brown  to  a 
usually  dark-brown  color.    Length,  5  to  7  mm. 

Form.  Head  distinctly  narrower  than  the  pronotum.  Vertex  usually 
distinctly  produced  medially,  about  half  longer  at  the  middle  than  next  the 
eyes  and  about  twice  as  broad  as  long.  Front  broad,  lorae  large,  cheeks  wide, 
clypeus  widened  apically.  Pronotum  over  twice  as  wide  as  long,  anterior 
margin  usually  a  little  more  curved  than  posterior,  the  lateral  and  humeral 
margins  subequal,  the  disc  transversely  wrinkled.  Scutellum  large,  the  surface 
granular.  Tegmina  moderately  long,  the  costal  margin  running  straight 
clear  to  the  apex,  forming  an  acute  tip. 

Color.  Vertex,  pronotum  and  scutellum  brownish  to  olive  green,  the 
scutellum  with  basal  angles  and  three  longitudinal  lines,  light.  Tegmina  usually 
shining  dark  brown,  sometimes  lighter.  When  dark  brown  the  nervures  are 
lighter  and  some  of  the  cells,  especially  along  the  costa  and  on  the  clavus,  are 
subhyaline  or  greenish.  Light  specimens  have  the  viens,  especially  apically, 
bordered  with  brown.  Face  olive  green,  unmarked,  or  with  faint  arcs  on  the 
front.  Below  olive  green,  marked  more  or  Ipss  with  dark  brown  or  black, 
especially  on  the  thorax,  coxse  and  tergites  of  the  abdomen. 

External  genitalia.  Female:  Last  ventral  segment  twice  as  long  as  preced- 
ing, broad  basally,  lateral  margins  rounding  to  slightly  notched  and  produced 
posterior  margin ;  pygofers  bearing  a  few  scattered  spines  and  slightly  exceeded 
by  ovipositor.  Male:  Valve  not  visible,  plates  long  and  narrow,  parallel- 
margined,  obtuse  apices  somewhat  divergent  and  greatly  exceeded  by  spiny 
P3'-gofers. 


118  The  University  Science  Bulletin. 

Internal  male  genitalia.  Styles  with  margins  of  anterior  half  sinuately 
tapering,  distal  half  stout  and  strongly  curved;  the  large,  club-shaped  and 
coarsely  granular  apices  strongly  diverging.  Connective  small,  heart-shaped, 
with  the  excision  wide  and  the  apex  broadly  rounding.  CEdagus  broad  basally, 
narrowing  to  the  middle,  bearing  two  small  basal  processes  and  a  larger  apical 
one  near  the  tip  of  which,  on  the  ventral  surface,  is  the  fimbriated  opening  of 
the  penis. 

Distribution.  This  species  is  largely  southern  and  eastern  in  its 
distribution.  The  many  specimens  examined  by  the  writer,  in  addi- 
tion to  those  mentioned  by  Van  Duzee,  are  distributed  as  follows: 
Charter  Oak,  Pa.  (J.  N.  KnuU) ;  Pt.  Royal,  Harrisburg,  Rockville, 
Pa.  (J.  G.  Sanders) ;  Lakehurst,  N.  J.  (J.  B.  Weiss) ;  Great  Falls, 
Md.,  Berkeley,  W.  Va.,  Ft.  Royal,  Va.,  Washington,  D.  C.  (Heide- 
mann) ;  Orangeburg,  S.  C.  (F.  H.  Lathrop) ;  Kansas  City,  Mo.  (F. 
Rogers) ;  Bisc  Bay,  Jacksonville,  Fla.,  Gainesville,  Fla.  (C.  J. 
Drake) ;  Ardmore,  Okla.  (F.  C.  Bishopp)  ;  Jacksonville,  Tex.  (W.  D. 
Pierce) ;  Boerne,  Tex.  (F.  C.  Pratt) ;  Victoria,  Rosser,  Tex.  (J.  D. 
Mitchell) ;  Kushla,  Ala.  (A.  H.  Sturtevant) ;  Alabama,  Florida, 
Mexico  (C.  F.  Baker)  ;  Knoxville,  Nashville,  Tenn.  (W.  B.  Cart- 
wright,  C.  C.  Hill) ;  Nashville,  Covey  Spring,  Chattanooga,  Tenn. 
(Geo.  G.  Ainslie) ;  Colliersville,  Clarksville,  Paris,  Lexington,  Tenn. 
(Dwight  M.  De  Long)  ;  Agricultural  College,  Mississippi  (H.  E. 
Weed) ;  Cherokee,  Bourbon  and  Miami  counties,  Kansas  (R.  H. 
Beamer) ;  Virginia,  labeled  Jassits  tructilis,  (Uhler)  ;  Spring  Creek, 
Okefinokee  Swamp,  Bainbridge,  Ga.  (J.  C.  Bradley) ;  Little  Rock, 
Ark.;  Capa,  S.  D. 

E.  L.  Dickerson  reports  what  are  presumably  this  species  from 
Cologne,  Lakehurst  and  Egg  Harbor,  N.  J. 

Remarks.  The  writer  has  examined  the  three  type  specimens,  one 
male  and  two  females,  of  Adnopterus  acuminatus  from  the  collec- 
tion of  the  Iowa  State  College,  and  the  female  type  from  the  Cornell 
LTniversity  collection.  The  two  female  specimens  from  Maryland 
and  Virginia  are  clearly  of  a  different  species  from  the  male  and  fe- 
male from  California,  as  shown  by  a  comparison  of  the  last  ventral 
segment  of  the  two  females.  It  seems  evident,  though,  that  the  ma- 
jority of  the  eight  specimens  from  which  Van  Duzee  described  the 
species  were  from  the  East  or  Southeast,  and  that  he  evidently  meant 
to  describe  a  species  with  such  a  distribution.  Accordingly  these 
eastern  females  are  retained  as  types  of  this  species,  while  the  two 
California  specimens,  along  with  a  large  amount  of  western  material, 
are  placed  in  the  following  species,  which  not  only  is  clearly  western 
in  its  distribution,  but  is  decidedly  different  as  to  the  genitalia  of 
both  males  and  females  from  the  above  species. 


Lawson:    Genus  AciNOPTERrs.  119 

Acinopterus  angulatus  n.  sp. 

(PI.  VIII,  fig.  8;   pi.  IX,  fig.  3;   pi.  X,  fig.  2;   pi.  XI,  fig.  4;   pi.  XII,  figs.  T),  6.) 

A  smaller  species  than  the  preceding,  vaiying  from  light"  to  dark  brown  in 
color,  but  uniformly  lighter  than  acuminatus  and  lacking  its  olive-green  tinge. 
Length,  5  to  6.25  mm. 

Form.  Head  broad,  scarcely  narrower  than  prothorax.  Vertex  broad  and 
short,  over  twice  as  wide  as  long,  about  one-third  longer  at  middle  than  next 
the  eyes,  anterior  margin  rounded  or  slightly  angulate,  sloping,  and  with  an 
impressed  line  behind  the  apex.  Front  broad  and  short,  lorse,  clypeus  and 
gense  characteristic  of  the  genus.  Pronotum  over  twice  as  wide  as  long,  an- 
terior margin  broadly  convex,  posterior  margin  shallowdy  concave,  lateral  and 
humeral  margins  distinct,  the  disc  transversely  wrinkled.  Scutellum  finely 
granulated,  the  transverse  impressed  line  curved  and  distinct.  Tegmina  with 
sutural  margin  rimning  straight  clear  to  the  tip,  forming  an  acute  apex. 
Venation  distinct,  with  from  one  to  several  cross-veins  between  the  first  and 
second  anal  veins. 

Color.  Vertex,  pronotum  and  scutellum  browTi  or  yellowish  brown.  Scu- 
tellum with  basal  angles  and  three  longitudinal  lines,  light.  Elytra  brown,  the 
veins  margined  with  darker  brown,  so  that  many  of  the  cells,  especially  along 
the  costa  and  apicallj^  appear  light.  Face  browTi;  front  with  eight  pairs  of 
dark-brown  lines,  wliich  are  longest  above  and  shortest  below,  leaving  the  mid- 
dle portion  unmarked.  Below  browm,  but  with  parts  of  thorax,  the  coxae  and 
femora  of  the  meso-  and  metathoracic  legs  and  the  dorsum  of  the  abdomen 
usuall}-  black  or  dark  brown. 

External  genitalia.  Female:  Last  ventral  segment  twice  as  long  as  pre- 
ceding, the  posterior  margin  varying  from  slightly  concave  to  slightly  convex, 
but  always  with  a  shght  median  notch  and  foiTning  very  distinct  lateral  angles 
with  the  long  lateral  margins.  Pygofers  moderately  wide,  sparsely  spined,  ex- 
ceeded shghtly  bj^  the  ovipositor.  Male:  Last  ventral  segment  long,  hiding 
the  valve.  Plates  broad,  about  the  length  of  the  last  ventral  segment,  only 
slight  h'  narrowing  to  the  obtuse  tips  which  are  divergent  medially,  but  have 
distinct  lateral  angles.  Pygofers  with  a  few  stout  spines  and  exceeding  the 
plates  by  about  two-thirds  the  length  of  the  latter. 

Internal  male  genitalia.  Styles  with  rather  small  and  pointed  anterior  ends, 
widest  at  point  of  the  distinct  process  to  the  connective,  then  strongly  curved 
to  the  very  wide  apices,  which  are  quite  granulated  and  clearly  concave  be- 
tween the  outer  angle  and  the  larger,  more  produced  inner  apex.  The  outer 
margin  bears  a  few  small  spines.  Connective  heart-shaped,  the  apex  quite 
broad.  (Edagus  of  the  pattern  characteristic  of  the  genus,  the  basal  processes 
well  developed  and  toothed,  the  terminal  process  veiy  long,  clearly  showing 
the  penis,  which  opens  at  the  fimbriate  extreme  apex  of  the  process. 

Distribution.  With  one  exception,  a  specimen  from  Washington, 
D.  C,  all  the  material  at  hand  came  from  the  South  and  West.  A 
single  specimen  was  taken  in  Yucatan,  many  specimens  coming 
from  other  parts  of  Mexico.  We  are  evidently  safe  in  calling  it  a 
southwestern  species,  which  may  possibly  extend  its  range  into  the 
southeastern  states.    The  following  are  the  locality  records  of  the 


120  The  University  Science  Bulletin. 

material  studied:  Riverside,  Chino,  San  Diego,  Ontario,  Visalia, 
Tia  Juana,  Oroville,  Caliente  (Ball)  ;  Hamilton  (no  collector's 
name)  ;  Los  Angeles  (Coquillett  and  A.  Koebele)  ;  Whittier,  Brawley 
(H.  0.  Osborn)  ;  all  from  California,  along  with  a  number  of  speci- 
mens taken  by  C.  F.  Baker;  Vera  Cruz  (no  collector's  name)  along 
with  two  specimens  taken  by  Gillette  and  a  series  by  Baker,  all  from 
Mexico;  Victoria  (J.  D.  Mitchell),  Orizaba  (H.  Osborn),  College 
Station  (no  collector's  name),  Brewster  county  (Mitchell  and  Cush- 
man),  all  from  Texas.  Also  a  single  specimen  taken  at  Progresso, 
Yucatan,  by  Frederick  Knab,  and  one  taken  at  Washington,  D.  C. 

Holotype,  male,  Ontario,  Cal.    Collection  of  E.  D.  Ball. 

Allotype,  female,  Ontario,  Cal.  Snow  collection,  University  of 
Kansas. 

Paratypes,  male,  Visalia,  Cal.,  and  female,  Chino,  Cal.,  in  col- 
lection of  E.  D.  Ball;  male,  Riverside,  Cal.,  and  female,  Chino,  Cal.. 
in  Snow  collection;  male,  Whittier,  Cal.,  and  female,  Brawley,  Cal., 
in  collection  of  H.  0.  Osborn. 

Acinopterus  inornatus  (Bak.) 

(PI.  VIII,  figs.  .3,  i   [type],  ,'5;   pi.  IX,  figs.  8,  9;   pi.  X.  fig.  (1;   pi.  XI,  fig.  7; 

pi.  XII,  figs.   11,   12.) 

Phlepsius  inornatus  Bak.,  Psyche,  vii,   Suppl.  i,  p.   13,   1895. 

Acinopterus  acuminatus  var.  inornatus  A'an  D.,  Cat.  Heniip.  N.  A.,  p.  675,  1917. 

The  following  is  the  original  description. 

Phlepsius  inornatus,  n.  sp.  Differing  from  all  other  species  of  the  genus 
in  being  entirely  destitute  of  elytral  reticulations  or  other  markings.  Length 
of  male,  6  mm. 

Male:  Head  narrower  than  the  pronotum.  Face  a  twelfth  wider  than  long; 
clypeus  one-half  longer  than  broad,  somewhat  constricted  before  the  base,  basal 
suture  strongly  curved,  apex  slightly  concave;  lorae  as  long  and  two-thirds  as 
broad  as  clypeus;  margin  of  gense  rather  slightly  incur\-ed  below  the  eye,  below 
this  strongly  conve.x,  thence  slightly  incurved  to  tip  of  clypeus.  Front  an 
eleventh  longer  than  broad,  somewhat  less  than  twice  the  length  of  the 
clypeus,  broad  below,  the  sides  very  slightly  incurved  at  the  antennse.  Disc 
of  the  vertex  flat,  length  at  middle  once  and  a  half  that  next  the  eye,  width 
between  the  eyes  once  and  a  half  the  length.  Width  of  the  pronotum  two  and 
a  third  times  the  length,  the  length  about  once  and  two-thirds  that  of  the 
vertex,  curvature  nearly  two-fifths  of  the  length,  posteriorly  irregularly  trans- 
versely wrinkled.  Scutellum  and  elytral  venation  normal.  Plate  not  visible, 
valves  two  and  a  half  times  longer  than  broad  at  base,  slightly  narrower  at 
apex,  blunt  at  tips,  without  hairs.  Pygofers  one-half  longer  than  valves, 
pointed  at  tips,  their  whole  outline  subtriangular,  provided  on  disc  of  lower 
surface  with  several  rather  long  whitish  spines  arranged  in  a  single  longitudinal 
row. 

Color  very  pale  yellowish,  deeper  on  the  abdomen.  Pronotum  with  five 
very  indistinct   longitudinal   whitish   bands.     Elytra  translucent,   pale   milky 


Lawson:    Genus  Acinopterus.  121 

white,  with  indistinct  smoky  clouds  on  the  discs  of  some  of  the  apical  and 
anteapical  areoles.  Veins  white,  claval  suture  brownish.  Face  and  legs  tinged 
with  greenish,  some  of  the  white  tibial  spines  brown  tipped.  Tarsal  joints  at 
apico;*  dark.  Dorsal  abdominal  segments,  fexcept  lateral  and  apical  margins, 
blackish. 

Described  from  a  single  male  taken  at  San  Augustine  (Ckll.  2140).  In  form 
this  insect  very  closely  resembles  P.  superbus  and  in  structure  is  strictly  con- 
generic with  it.  It  differs  very  widely,  however  (and  this  is  a  generic  differ- 
ence according  to  Van  Duzee's  sj'noptic  table  of  the  genera),  in  that  it  does  not 
possess  the  elytral  reticulations  or  other  markings  so  characteristic  of  the 
genus.  On  a  very  superficial  examination  it  might  be  taken  for  a  Chlorotettix, 
but  its  general  form,  stronger  build,  and  lengthened  vertex  are  strictly  Phlep- 
siid. 

The  writer  gives  the  following  description: 

A  rather  slender  and  light-colored  species  which  sometimes  may  be  rather 
dark.    Length,  5  to  6.5  mm. 

Form.  Head  distinctly  narrower  than  the  pronotum.  Vertex  varying  in 
length,  but  usually  about  twice  as  wide  as  long  and  half  longer  at  the  middle 
than  next  the  eye;  disc  sloping  and  with  the  impressed  line  behind  the  apex. 
Face  as  in  the  other  members  of  the  genus.  Pronotum  over  twice  as  wide 
as  long,  the  anterior  margin  more  strongly  curved  than  the  posterior,  the 
lateral  and  humeral  margins  about  equal.  Scutellum  of  average  size,  finely 
granulated,  and  with  distinct  transverse  impressed  line.  Tegmina  long  and 
narrow,  sutural  margin  extending  straight  clear  to  tip,  forming  an  acute 
apex;  venation  usually  distinct,  though  sometimes  rather  weak,  with  from  one 
to  several  cross-veins  between  the  first  and  second  anal  veins. 

Color.  Yellowish  or  yellowish  grey.  Pronotum  with  five  pale  and  some- 
times indistinct  lines.  The  tegmina  vary  considerably,  being  sometimes 
almost  colorless  till  near  the  tip,  while  in  others  the  veins  are  margined  lightly 
with  brown,  especially  at  the  margins,  but  in  all  cases  some  of  the  apical  cells 
are  more  or  less  darkened.  The  darkened  tips  of  the  veins  along  the  sutural 
and  costal  margins  sometimes  give  the  elytra  a  variegated  appearance.  Be- 
neath this  species  is  usually  light  except  for  the  darkened  doreum  of  the 
abdomen. 

External  genitalia.  Female:  Last  ventral  segment  over  twice  as  long  as  the 
preceding,  posterior  margin  di.stinctly  but  roundingly  produced  medially,  but 
with  the  small  median  notch  characteristic  of  the  genus.  Pygofers  moderately 
wide,  sparsely  spined,  and  slightly  exceeded  by  the  ovipositor.  Male:  Last 
ventral  segment  longer  than  the  preceding,  hiding  the  valve.  Plates  long  and 
fingerlike,  about  the  length  of  the  last  ventral  segment,  shghtly  narrowed  to 
the  somewhat  diverging  but  rounded  tips.  Pj'gofers  bearing  a  few  stout  spines 
and  exceeding  the  plates  by  about  two-thirds  the  length  of  the  latter. 

Internal  male  genitalia.  Styles  wide  basally  and  with  strong  process  to  con- 
nective; apically  strongly  curv^ed  and  clublike,  the  apical  portion  of  the  club 
with  distinct  and  large  granulations,  giving  it  a  rough  appearance.  Connective 
heart-shaped,  the  excision  wide  and  shallow,  the  apex  rounded,  ffidagus  rather 
small  but  stout,  the  upper  part  much  like  an  inverted  boot,  the  heel  distinctly 
cleft ;  the  paired  basal  processes  about  half  the  length  of  the  apical  process, 
their  lower  edges  serrate;  the  terminal  process  stout,  the  penis  opening  at  the 
fimbriate  tip. 


122  The  University  Science  Bulletin. 

Distribution.  This  species  is  evidently  a  southwestern  form,  for 
all  the  material  at  hand  was  taken  in  this  region.  The  following 
are  the  locality  records  of  the  twenty-two  specimens  examined: 
Type  specimens  from  San  Augustine,  N.  M.  (C.  F.  Baker) ;  On- 
tario, Riverside,  Cal.  (E.  D.  Ball)  ;  Santa  Rita  mountains  (F.  H. 
Snow),  Galiuro  mountains  (H.  G.  Hubbard),  Phoenix  (E.  D.  Ball), 
Sabino  Canyon,  St.  Catalina  mountains  (E.  L.  Dickerson) ,  all  from 
Arizona;  Brewster  county,  Texas  (Mitchell  and  Cushman). 

Remarks.  As  far  as  the  writer  knows,  this  species  has  been  known 
hitherto  only  from  the  type.  In  the  material  gathered  for  the  study 
of  the  genus  he  found  a  number  of  similar  specimens,  which,  while 
differing  in  some  ways,  particularly  in  the  length  of  the  vertex  and 
the  extent  of  the  elytral  markings,  are  yet  thought  to  be  representa- 
tives of  this  species,  for  a  careful  study  of  the  genitalia  of  several 
males  revealed  no  differences,  although  the  vertices  of  the  specimens 
were  quite  unlike.  Also  the  specimens  show  a  complete  range  in 
color  from  that  of  the  very  light  type  specimen  to  forms  that  are 
distinctly  brownish.  While  frankly  having  some  doubt  as  to  the 
specific  identity  of  all  the  material  named  thus,  the  writer  feels  it 
better  to  call  them  all  the  same  species  rather  than  to  describe  new 
species  on  insufficient  material. 

Acinopterus  productus  n.  sp. 

(PI.  VIII,  fig.  G;   pi.  IX,  fig.  2;   pi.  X,  fig.  3;   pi.  XI,  fig.  3;   pi.  XII,  figs.   13,  14.) 

A  distinctly  greenish  species,  differing  from  other  green  forms  by  the  acute 
apex  of  the  tegmina  and  the  produced  vertex. 

Form.  Head  distinctly  narrower  than  pronotum.  Vertex  less  than  twice  as 
wide  as  long,  at  least  half  longer  at  the  middle  than  next  the  eye,  the  apex 
rounded  and  with  an  impressed  line  parallel  with  the  margin.  Face  character- 
istic of  the  genus.  Pronotum  short,  well  over  twice  as  wide  as  long,  the  ante- 
rior margin  more  strongly  curved  than  the  posterior,  the  lateral  and  humeral 
margins  about  equal,  the  disc  transversely  wrinkled.  Scutellum  with  the  usual 
granular  surface  8,nd  impressed  line.  Tegmina  long  and  rather  narrow,  the 
costal  margin  running  straight  clear  to  the  tip,  forming  an  acute  apex,  and 
with  the  nervures  distinct,  the  fii-st  and  second  anal  veins  usually  united  by 
several  cross-veins. 

Color.  The  entire  insect  is  green  except  for  the  darkened  apices  of  the  ely- 
tra and  the  yellowish  or  pinkish  legs.  The  nervures  stand  out  as  a  lighter 
green  than  the  cells  of  the  tegmina. 

External  genitalia.  Female:  Last  ventral  segment  over  twice  as  long  as 
the  preceding,  the  medially  produced  posterior  margin  with  the  usual  small 
notch,  the  lateral  margins  rounding  with  the  posterior.  Pygofers  rather  ro- 
bust, sparsely  spined,  and  slightly  exceeded  by  the  ovipositor.  Male:  Valve 
hidden  by  the  long  last  ventral  segment.  Plates  longer  than  last  ventral  seg- 
ment, fairly  wide  basally,  tapering  to  the  divergent  and  rounded  but  com- 


Lawson:    Genus  Acinopterus.  123 

parativeh'  narrow  ajiex.    Sparsely  bristled  pygofers  exceeding  i)]a1es  by  about 
two-thirds  the  length  of  the  latter. 

Internal  male,  (icnitalia.  Styles  of  usual  form,  widest  at  point  of  process  to 
connective,  apically  club-shaped,  the  granulated  and  blunt  apices  slightly  but 
clearly  concave.  Connective  heart-shaped,  the  excision  fairly  deep,  ffidagus 
with  body  as  in  other  species,  the  basal  processes  small,  the  terminal  process 
of  medium  diameter  and  length. 

Distribution.  The  eight  specimens  at  hand  when  this  species  was 
described  all  came  from  California  and  Arizona.  They  were  all 
taken  by  Dr.  E.  D.  Ball.  The  California  specimens  are  from  Im- 
perial, Beaumont  and  Riverside,  while  the  two  Arizona  specimens 
are  from  Phoenix. 

Holotype,  female,  Imperial,  Cal..  in  collection  of  Doctor  Ball. 

Allotype,  male,  and  paratype,  female,  both  from  Imperial,  Cal., 
in  the  Snow  collection,  University  of  Kansas. 

Paratypes,  male  from  Imperial  and  female  from  Beamnont,  Cal., 
in  collection  of  Doctor  Ball. 

Acinopterus  brunneus  Ball. 

(PI.  VIII,  fig.  2;   pi.  IX,  fig.  5;   pi.  X,  fig.  7;    pi.  XI,  fig.  6;    pi.  XII,  figs.   9,  10.) 

Acinopterus  acuminatus  var.   brunneus  Ball,  Can.  Ent.,  xxxv,  p.  231,   1903. 
Acinopterus  acuminatus  var.  brunneus  Van  D.,  Cat.  Hemip.  N.  A.,  p.  675,  1917. 

The  following  is  the  original  description : 

^4.  acuminatiifi,  var.  brunncuf;,  n.  var.  Shghtly  larger  than  the  preceding 
variety.  "\'ertex,  pronotum  and  scutellum  pale  green,  washed  with  cinnamon 
brown.  Elytra  pale  cinnamon  brown,  slightly  fuscous  at  tip.  Whole  insect 
with  a  slight  tawny  iridescence,  below  pale  green. 

Described  from  three  specimens  from  Rifle,  Colo.,  taken  by  the  author. 

• 
The  writer  adds  the  following  description: 

A  large  brownish  or  greenish-brown  species,  about  the  largest  member  of 
the  genus.    Length,  5.5  to  6.75  mm. 

Form.  Head  about  as  wide  as  the  pronotum.  Vertex  at  least  twice  as  wide 
as  long,  one-third  longer  at  middle  than  next  the  eye,  the  anterior  margin 
rounded,  and  with  the  characteristic  depression  behind  the  apex.  Face  with 
all  the  parts  ver\'  broad,  the  lorse  nearly  reaching  the  margin  of  the  genae. 
Pronotum  over  twice  as  wide  as  long,  the  anterior  margin  more  strongly  cur\^ed 
than  the  posterior,  lateral  and  humeral  margins  distinct  and  about  equal,  the 
disc  transversely  WTinkled.  Scutellum  as  in  other  members  of  the  genus.  Teg- 
mina  with  rounded  apex  but  more  acute  than  in  viridis,  the  venation  often  less 
distinct  than  in  other  species,  and  usually  with  several  cross-veins  between  the 
first  and  second  anal  veins. 

Color.  Vertex,  pronotum  and  scutellum  greenish-brown.  Tegmina  of  same 
color  or  darker,  the  veins  of  the  apical  half  often  being  margined  with  dark 
brown,  giving  the  tip  a  darker  appearance.  Beneath  the  color  is  usually  as 
above  but  sometimes  the  hind  legs  and  abdomen  have  a  reddish  tinge. 


124  The  University  Science  Bulletin. 

External  genitalia.  Female :  Last  ventral  segment  differing  from  that  of 
any  other  member  of  the  genus  in  being  extremely  produced  medially,  three 
times  as  long  as  the  preceding  segment,  with  the  usual  small  apical  excision, 
and  the  lateral  margins  sometimes  slightly  concave.  The  broad  and  spiny 
pygofers  are  slightly  exceeded  by  the  ovipositor.  Male:  Last  ventral  segment 
long,  hiding  the  valve.  Plates  long  and  slender,  longer  than  last  ventral  seg- 
ment, their  sides  straight  to  the  somewhat  narrowed  but  rounded  apices. 
Pygofers  long,  exceeding  plates  by  about  the  length  of  the  latter,  bearing  the 
usual  spines. 

Internal  male  genitalia.  Styles  stout,  apical  part  nearly  of  same  width 
throughout  and  bearing  many  granulations.  Connective  as  broad  apically  as 
basally.  CEdagus  unhke  anything  in  the  genus  and  very  characteristic,  the 
basal  processes  short  and  not  serrate,  the  distal  process  verj'  long,  the  fimbriate 
opening  of  the  penis  extending  back  from  the  extreme  tip  for  a  considerable 
distance.  In  addition  a  pair  of  large  ventral  and  lateral  lobes  is  present  that 
completely  cover  the  basal  processes,  these  lobes  being  serrate  along  their  ven- 
tral margin.  The  entire  dorsal  part  of  the  oedagus  is  also  quite  different  from 
the  corresponding  part  in  the  other  species  of  the  genus. 

Distribution.  With  the  exception  of  a  single  specimen  taken  by 
Coquillett  at  Los  Angeles,  Cal.,  all  the  other  fourteen  specimens  ex- 
amined by  the  writer  are  from  Doctor  Ball's  collection.  These  were 
taken  from  the  following  localities:  Rifle,  Colo.;  Pardman,  Salt 
Lake  City,  Utah;  Ravenna,  Cabazon,  Riverside,  and  Beamnont,  Cal. 

Acinopterus  pallidus  n.  sp. 

(PI.  Vlll,  fig.  10;   pi.  IX,  fig.  4;   pi.  XI,  fig.  2.) 

Closely  related  to  the  preceding  species,  but  slightly  smaller  and  paler. 
Length,  5.5  to  6  mm. 

Form.  Head  distinctly  narrower  than  the  pronotum.  Vertex  a  little  over 
twice  as  wide  as  long,  not  quite  one-third  longer  at  the  middle  than  next  the 
eye,  the  anterior  margin  evenly  rounded  and  broadly  rounding  with  the  front, 
the  depression  back  of  the  apex  small.  Face  very  broad,  the  front  fully  as 
broad  basally  as  long,  and  the  gense  quite  wide.  Pronotum  over  twice  as  wide 
as  long,  anterior  margin  but  slightly  more  curved  than  the  posterior,  lateral 
and  humeral  margins  about  equal,  disc  with  transverse  wrinkles.  Scutelkun 
with  usual  granular  surface  and  transverse  impressed  line.  Tegmina  moder- 
ately long,  the  apices  more  rounded  than  in  preceding  species,  venation  dis- 
tinct but  not  conspicuous,  and  with  but  one  or  two  cross-veins  between  the 
first  and  second  anal  veins. 

Color.  The  entire  insect,  abov^e  and  below,  of  a  pale  greenish-yellow  color. 
Apices  of  some  of  the  veins  along  costal  margin  and  at  apex  margined  with 
black,  making  the  tips  of  the  tegmina  appear  slightly  darkened.  The  tarsi 
tend  to  be  brownish. 

External  genitalia.  Female :  Last  ventral  segment  produced  medially,  bear- 
ing a  shallow  median  notch  apically  from  which  the  margins  round  to  the 
base.    Pygofers  large,  sparsely  spined,  exceeded  by  the  ovipositor. 

Distribution.  Described  from  four  specimens  taken  by  Doctor 
Ball  at  Cabazon,  Cal. 


Lawson:    Genus  Acinopterus.  125 

Holotype,  female,  and  two  paratypes,  females,  in  collection  of 
Doctor  Ball. 

One  paratype,  female,  in  the  Snow  collection,  University  of 
Kansas. 

Remarks.  This  species  stands  between  brunneus  and  viridis. 
From  the  former  it  differs  in  its  smaller  size,  lighter  color,  and 
shorter  female  ventral  segment,  while  from  the  latter  it  differs  in 
being  lighter,  and  in  not  having  the  tegminal  veins  green. 

Acinopterus  viridis  Ball. 

(PI.  VIII,  fig.  11;  pi.  IX,  fig.  6;  pi.  X,  fig.  4;  pi.  XI,  fig.  5;  pi.  XII,  figs.  7,  8.) 

Acinopterus  acuminatus  var.  viridis  Ball,  Can.  Ent.,  xxxv,  p.  231,  1903. 
Acinopterus  acuminatus  var.  viridis  Van  D.,  Cat.  Hemip.  N.  A.,  p.  675,  1917. 
Acinopterus  acrimirtatus  var.  viridis  Laws.,  Kan.   Univ.  Sci.  Bui.,  xii,  p.  208,  1920. 

The  following  is  the  original  description: 

A.  acuminatus,  var.  viridis,  n.  var.  Form  and  structure  of  the  preceding 
nearly;  slighth-  smaller.  Bright  grass  green  both  above  and  below.  Eyes 
and  extreme  tip  of  elytra  fuscous. 

Described  from  a  number  of  specimens  from  southern  Colorado  and  Ari- 
zona. This  is  the  common  form  in  southern  Colorado,  where  it  was  taken  by 
E.  P.  Van  Duzee  and  the  author. 

The  following  description  is  by  the  writer : 

A  rather  robust  greenish  species,  with  or  without  elytral  markings.  Length, 
5  to  6  mm. 

Form.  Head  distinctly  narrower  than  the  pronotum.  Vertex  about  twice 
as  wide  as  long,  one-half  longer  at  the  middle  than  next  the  eye,  anterior 
margin  broadly  rounded  and  with  a  depression  just  behind  apex.  Face  with 
all  the  sclerites  broad.  Pronotum  over  twice  as  wide  as  long,  anterior  margin 
more  strongly  curved  than  the  posterior,  lateral  and  humeral  margins  about 
equal,  the  disc  transversely  wrinkled.  Scutellum  as  in  other  members  of  the 
genus.  Tegmina  moderately  long,  the  apex  narrowed  but  rounded,  with 
usually  one  or  two  or  sometimes  several  cross-veins  between  the  first  and 
second  anal  veins  and  sometimes  one  or  two  between  the  second  and  third. 

Color.  Vertex,  pronotum  and  scutellum  usuall}^  green,  though  sometimes 
distinctly  j'ellowish.  Tegmina  green  with  the  nervures  darker  green,  the 
latter  being  sometimes  not  margined  at  all  or  bordered  with  brown  till  all 
the  apices  of  the  veins  at  the  sutural  margin,  along  the  distal  half  of  the 
costal  margin,  and  at  the  apex,  are  definitely  bordered,  frequently  giving  the 
apex  a  darker  appearance.  Below  the  insect  is  also  green,  the  tarsi  tending 
to  be  brownish. 

External  genitalia.  Female :  Last  ventral  segment  over  twice  as  long  as 
the  preceding,  the  lateral  and  posterior  margins  rounding  to  the  produced  apex 
which  bears  the  usual  small  median  notch.  Pygofers  stout,  sparsely  spined, 
exceeded  by  the  ovipositor.  Male:  Last  ventral  segment  long,  hiding  the 
valve.  Plates  long  and  slender,  slightly  longer  than  last  ventral  segment,  the 
bases  distinctlj'  wider  than  the  divergent  apices.  Pygofers  broad,  sparsely 
spined,  exceeding  the  plates  by  about  two-thirds  the  length  of  the  latter. 

Internal  male  genitalia.    Stjdes  large,  widest  at  point  of  process  to  connec- 


126  The  University  Science  Bulletin. 

tive,  distal  portion  strongly  curved  and  then  running  straight  to  the  expanded 
tips,  which  have  the  inner  angles  about  right-angled,  but  the  outer  angles 
strongly  produced,  the  distal  margin  between  the  two  corners  being  slightly  con- 
cave. The  outer  margins  of  the  distal  half  are  roughened  and  the  characteris- 
tic granulations  appear  over  the  entire  apical  portion.  Connective  heart-shaped, 
the  apex  broadly  rounded,  ffidagus  very  characteristic  of  the  species,  having 
two  pairs  of  basal  processes,  the  upper  ones  shorter,  the  lower  ones  reaching 
nearly  to  the  tip  of  the  apical  process,  at  the  extreme  fimbriate  tip  of  which 
the  penis  opens.    Both  pairs  of  basal  processes  bear  teeth  along  the  margins. 

Distribution.  All  of  the  twenty-five  specimens,  except  one  from 
Colorado,  one  from  Morton  county,  Kansas,  and  one  from  Ashfork, 
Ariz.  (Barber  and  Schwarz),  were  sent  the  writer  by  Doctor  Ball, 
who  obtained  them  from  the  following  localities:  Soldier,  Dixie, 
Richfield,  Monroe,  Moab,  Utah;  Fort  Collins,  Grand  Junction, 
Delta,  Dutch  George  or  Poudre  Canyon,  Colo.;  Coolidge,  Kan.; 
Wenatchee,  Wash. ;  Phoenix,  Ariz.  There  are  specimens  also  in  the 
collection  of  the  Colorado  Agricultural  College  from  some  of  these 
localities. 

Remarks.  The  specimen  from  Moab,  Utah,  seems  to  be  different 
from  the  other  specimens  of  this  species  in  that  it  is  lighter  in  color, 
has  a  broader  head,  and  longer  and  more  pointed  elytra.  There  be- 
ing only  one  specimen  of  its  kind,  however,  the  writer  prefers  to 
place  it  here  to  describing  it  as  a  new  species  from  a  single  speci- 
men. 

Acinopterus  viridis  var.  variegatus  Ball. 

(PI.  VIIl,  fig.  7 ;   pi.  IX,  fig.  7.) 

Acmopterus  acuminatus  var.   variegatus  Ball,  Can.  Ent.,  xxxv,  p.  231,   1903. 
Acinopterus  acuminatus  var.  inornatus  Van.  D.,  Cat.  Hemip.  N.  A.,  p.   675,  1917. 

The  following  is  the  original  description: 

Acinopterus  acuminatus,  var.  variegatus,  n.  var.  Form  and  structure  of  the 
species,  but  much  lighter  colored.  Vertex,  pronotum  and  scutellum  inclined 
to  be  reddish,  especially  in  the  male.  Elytra  whitish  pruinose,  nervures  green- 
ish, not  margined,  except  towards  apex  and  along  sutural  margin,  three  fuscous 
points  along  the  suture,  and  sometimes  one  on  the  disc  of  each  elytron. 

Described  from  twenty-four  specimens  from  Colorado  and  Arizona. 

The  following  is  the  writer's  description: 

The  members  of  this  variety  are  hke  viridis  except  in  color. 

Color.  General  color,  brown.  Vertex,  pronotum  and  scutellum  greyish  or 
brownish,  sometimes  with  a  reddish  tinge.  Tegmina  pale,  but  with  nervures 
margined  more  or  less  throughout,  especially  along  sutural  and  costal  margins 
and  apically,  giving  them  a  decidedly  variegated  appearance. 

Distribution.  Eight  specimens  examined  are  from  Doctor  Ball's 
collection  and  were  taken  by  him  at  Fort  Collins  and  Denver,  Colo. 


Lawson:    Genus  Acinopterus.  127 

In  the  collection  of  the  Colorado  Agricultural  College  are  other 
specimens  from  the  same  localities. 

Remarks.  The  specimens  at  hand  show  gradual  gradations  into 
the  variegated  form  of  viridis,  which,  in  its  turn,  goes  by  insensible 
gradations  into  the  pure  green  form  characteristic  of  the  species.  An 
examination  of  the  male  internal  genitalia  of  typical  variegatus  and 
that  of  a  variegated  viridis  showed  no  differences,  and  the  gradual 
loss  of  the  tegminal  markings  into  the  plain  green  form  would  seem 
to  indicate  the  identity  of  these  two  green  forms. 

Acinopterus  obtutus  n.  sp. 

(PI.  VIII,  fig.  9;   pi.  X,  fig.  5;   pi.  XII,  fig.s.  3,  4.) 

A  rather  small  green  species  with  a  relatively  larger  vertex  than  viridis. 
Length.  5.5  mm. 

Form.  Head  di.stinctl.>-  narrower  than  pronotiim.  Vertex  large,  about  twice 
as  wide  as  long,  one-half  longer  at  the  middle  than  next  the  eye,  a  sHght  de- 
pression just  behind  the  broadly  rounded  apex  which  rounds  very  obtusely  with 
the  front.  All  the  sclerites  of  the  face  rather  broad.  Pronotum  over  twice  as 
wide  as  long,  the  anterior  margin  more  curved  than  the  posterior,  the  lateral 
and  humeral  margins  about  equal,  the  transverse  wrinkles  of  the  disc  indis- 
tinct. Scutellum  with  the  usual  granulated  surface  and  transverse  impressed 
line.  Tegmina  with  the  tips  broken  off  in  all  three  of  the  specimens  from 
which  the  species  is  described,  but  presumably  rather  rounded  apically,  judging 
from  the  material  studied.  Claval  area  with  a  few  cross-veins  between  the 
first  and  second  anal  veins. 

Color.  Vertex,  pronotum  and  scutellum  green,  the  scutellum  with  basal 
angles  and  three  longitudinal  lines,  light.  Tegmina  milky  green,  the  veins 
light  or  dark  green,  and  margined  more  or  less  with  brown.  Below  the  entire 
insect  is  green. 

External  genitalia.  Male:  Last  ventral  segment  long,  hiding  the  valve. 
Plates  long  and  narrow,  nearly  parallel-margined  to  the  somewhat  divergent 
apices,  which  are  exceeded  by  the  bristly  pygofers  by  about  two-thirds  the 
length  of  the  former. 

Internal  male  genitalia.  Styles  of  the  usual  shape,  the  granular  apical  por- 
tions quite  expanded  at  the  tip,  which  is  either  straight  or  slightly  concave  at 
the  end.  Connective  nearly  round,  the  basal  excision  rather  deep.  Qildagus 
differing  from  that  of  any  member  of  the  genus.  The  body  is  very  wide 
ba.sally,  the  heel  cleft,  then  strongly  narrowed  to  the  base  of  the  procesvses,  of 
which  the  basal  pa'ir  are  short  and  slender  while  the  distal  one  is  quite  long 
and  slender,  the  penis  opening  at  the  extreme  fimbriate  tip. 

Distribution.  The  three  male  specimens  from  which  this  species 
is  described  are  all  from  the  collection  of  Mr.  E.  L.  Dickerson  and 
were  taken  in  the  Sabino  canyon  of  the  St,  Catalina  mountains  of 
Arizona. 

Remarks.  This  species  is  decidedly  smaller  than  the  other  green 
species,  and  the  cedagus  is  so  characteristic  that  on  this  structure 


128  The  University  Science  Bulletin. 

alone  the  writer  is  confidently  basing  the  distinctness  of  thefi6  spe- 
cies. Among  the  specimens  of  viridis  from  Colorado  and  Utah  there 
may  be  one  or  two  that  are  the  females  of  this  species,  but  not  being 
as  sure  of  their  position  as  of  these  three  males,  they  are  left  in  the 
former  group. 

Holotype  in  collection  of  Mr.  E.  L.  Dickerson. 

Paratypes  in  collection  of  Doctor  Ball  and  the  Snow  Collection. 
University  of  Kansas. 


PLATE  VIII. 

Fig.  1.   ..4.  acuminatus. 

Fig.  2.    A.  brunneus. 

Figs.  3,  4,  5.   A.  inornatus. 

Fig.  6.    A.  productus. 

Fig.  7.   A.  viridis  var.  variegatus. 

Fig.  8.   A.  angulalus. 

Fig.  9.   A.  obtutits. 

Fig.  10.   A.  pallidus. 

Fig.  11.    A.  viridis. 

(130) 


Lawson:    Genus  Acinopterxs. 


131 


PLATE  VIII. 


PLATE  IX. 

Fig.  1.  A.  acuminatus. 

Fig.  2.  A.  productus. 

Fig.  3.  A.  angulatus. 

Fig.  4.  A.  pallidus. 

Fig.  5.  A.  hrunneus. 

Fig.  6.  A.  viridis. 

Fig.  7.  ^.  viridis  var.  variegatus. 

Figs.  8  and  9.    A.  inornatus. 


(132) 


Lawson:    Genus  Acinopterus. 


133 


PLATE  IX. 


PLATE  X. 

Fig.  1.  A.  acuminatus. 

Fig.  2.  A.  angulatus. 

Fig.  3.  A.  productus. 

Fig.  4.  A.  viridis. 

Fig.  5.  A.  ^midm.  a^^J^iy<o 

Fig.  6.  A.  inornatus. 

V](i.  7.  A.  brunneus. 

(134) 


Lawson:    Genus  Acinopterus. 


135 


PLATE  X. 


PLATE  XI. 

Fig.  1. 

A.  acuminatus 

Fig.  2. 

A.  pallidus. 

Fig.  3. 

A.  productiis. 

Fig.  4. 

A.  angulatus. 

Fig.  5. 

A.  viridis. 

Fig.  6. 

A.  brunneus. 

Fig.  7. 

A.  inornatus. 

(136) 


Lawson:    Genus  Acinopterls. 


137 


PLATE  XI. 


PLATE  XII. 

Fitis.  1  AND  2.   A.  acuminatus. 
Figs.  3  and  4.   A.  ohlulus. 
Figs.  5  and  6.   A.  angulatus. 
Figs.  7  and  8.   A.  viridis. 
Figs.  9  and  10.   A.  brunneus. 
Figs.  11  and  12.   A.  inornatus. 
Figs.  13  and  14.   A.  productus. 

(138) 


J 


Lawson:    Genus  Acinopterus. 


139 


PLATE  XII. 


THE 

KANSAS  UNIVEKSITY 

Science  Bulletin 


Vol.  XIV,  No.  5— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  5.) 

ENTOMOLOGY  NUMBER  V. 


.    CONTENTS: 

The  Life  History  of  the  Toad  Bug  (Heteroptera), 

H.  B.  Hunger j or d. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  oflSce  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVEBSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  5. 


The  Life  History  of  the  Toad  Bug. 

Gelastocoris  oculatus  Fabr.  (Gelastocoridse).* 

BY  H.  B.  HUNGERFORD, 

Professor  of  Entomology,  University  of  Kansas. 

INTRODUCTION. 

IN  MY  PAPER  on  "The  Biology  and  Ecology  of  the  Aquatic 
Hemiptera"t  (pages  49-51)  I  gave  the  gist  of  what  was  known  at 
the  time  concerning  the  habits  and  life  history  of  the  toad  bug.  Its 
habitat  and  feeding  habits,  together  with  a  brief  description  of  the 
ovum  and  fifth  nymphal  instar  were  given  there. 

During  the  season  of  1920  I  had  an  opportunity  to  gather  con- 
siderable data  relative  to  these  interesting  insects.  Mrs.  Grace 
Wiley,  a  student  of  mine,  knpwing  of  my  desire  to  study  Gelasto- 
coris,  sent  me  a  number  of  living  adults  from  her  home  in  Chanute, 
Kan.,  in  the  autumn  of  1919.  One  of  these  I  kept  alive  until  Sep- 
tember, 1920.  On  May  14,  1920,  she  sent  me  a  shipment  of  adults, 
and  again  on  July  6  another  small  lot  of  the  bugs.  The  live  insects 
supplied  by  Mrs.  Wiley  thus  made  possible  the  notes  here  reported, 
and  I  wish  to  acknowledge  my  gratitude  to  her  for  her  kindness. 

THE  TECHNIQUE  USED  IN  THE  REARINGS. 

Tall  stenders,  or  staining  jars,  of  glass  about  the  size  of  jelly 
glasses  were  used.  In  each  of  these  was  placed  an  inch  of  sand 
that  had  been  sterilized  by  heat.  The  paired  adults  were  confined 
in  low  stenders  of  various  sizes,  and  the  sand  searched  each  day  for 
eggs.  The  young  were  isolated  in  the  tall  stenders  as  soon  as 
hatched,  for  they  were  cannibalistically  inclined,  and  two  young 

*  Mr.  Bueno  recognizes  this  as  a  new  species,  G.  acciduiis  Ms.  I  confess  I  cannot  dis- 
tinguish it  from  G.  oculatus  Fabr. 

t  Kansas  University  Science  Bulletin,  vol.  XI,  Dec.   1919;    265   pages,  33  plates. 

(145) 


146  The  University  Science  Bulletin. 

would  get  on  but  a  few  hours  together.  The  sand  was  moistened 
each  day,  and  the  jars  covered  with  ground-glass  covers.  Each 
nymph  was  examined  daily  for  molts,  which  were  removed  and 
placed  in  vials  of  alcohol  or  on  cotton  in  small  tin  boxes,  and  each 
instar  skin  of  each  insect  reared  was  preserved  separately  for  study. 
This  has  provided  adequate  material  for  study  of  structural  details 
of  each  stage. 

We  endeavored  to  determine  whether  the  insects  had  any  choice 
of  soil,  by  placing  them  in  pans  containing  sand  on  one  side  and 
sandy  loam  on  the  other.  Our  results  were  not  conclusive.  We 
also  used  sterilized  sandy  loam  in  some  of  the  rearing  jars  instead 
of  the  sand  and  found  it  of  no  advantage. 

The  insects  were  fed  house  flies,  plant  lice,  oscinid  flies,  cicadellids 
and  many  other  small  insects  taken  in  sweeping  the  grass  upon  the 
campus.  Each  day  the  dead  carcasses  were  cleaned  out  of  the  rear- 
ing jars  and  freshly  killed  insects  inserted. 

Mortality  was  very  high,  as  a  glance  at  the  tables  presented  be- 
low will  show,  and  indicates  that  some  essential  factor  of  their 
natural  habitat  was  lacking.  The  fact  that  116  nymphs  out  of  179 
died  in  the  first  stage,  and  that  they  usually  succumbed  on  the 
date  when  molting  might  have  been  expected  to  occur,  would  point 
to  a  hazard  of  ecdysis.  An  examination  of  the  dates  of  death  of 
the  older  nymphs  further  substantiates  this  view. 

Toward  the  end  of  a  stadium  the  nymph  always  appears  plump 
instead  of  flat,  and  so  the  appearance  of  the  nymph  indicates  ap- 
proximately its  development.  Thus  some  would  become  plump  in 
a  week  and  molt;  others  would  develop  more  slowly. 

In  spite  of  the  discouragements  because  of  the  very  high  rate  of 
mortality,  the  tending  and  study  of  the  rearings  were  very  interest- 
ing. Mr.  William  Hoffman,  who  assisted  me  very  materially  dur- 
ing the  latter  part  of  the  summer,  found  the  task  most  absorbing. 
He  fed  the  nymphs  and  kept  the  records  with  as  much  interest, 
care  and  ingenuity  as  I  could  have  done,  and  I  desire  to  acknowledge 
herewith  my  indebtedness  to  him  for  his  services. 

HABITAT. 

The  toad  bug  is  a  shore  bug,  found  along  the  muddy  banks  of 
small  streams  or  the  sandy  beaches  by  the  river.  It  is  a  notable 
example  of  protective  coloration.  Specimens  taken  on  muddy  banks 
are  dull  and  slaty  grey  with  indistinct  pattern,  while  those  from 
the  sandy  beaches  are  variegated,  pebbled  and  mottled  like  the 
sand.     Specimens  in  captivity  have  been  observed  to  burrow  be- 


Hungerford:    The  Toad  Bug.  147 

neath  tlie  dirt  in  cloudy  weather  and  remain  tlius  many  liours.  This 
habit  may  be  their  method  of  maintaining  their  geographical  posi- 
tion in  time  of  flood.  I  have  observed  broad,  sandy,  barren  flats 
where  toad  bugs  lived  become  inundated  by  rapid  currents  of 
water  for  a  few  hours;  nevertheless,  when  the  water  receded  and 
the  sun  came  again,  here  were  the  bugs  as  before.  There  were 
neither  vegetation  nor  sizable  stones  for  their  anchorage,  so  I  sup- 
pose they  "dug  in."  They  hop  about  with  considerable  alacrity 
when  alarmed  (one  first  instar  nymph  jumped  ten  inches).  They 
pounce  upon  their  prey,  which  appears  to  consist  of  almost  any 
sort  of .  insect  they  can  capture,  from  a  grouse  locust  (tettix)  to  a 
lacebug. 

MATING. 

There  is  considerable  sexual  dimorphism  with  the  toad  bug.  The 
abdomen  of  the  male  is  strongly  asymmetrical,  as  is  also  the  case  in 
the  male  Corixidse.  Figures  illustrating  this  are  shown  on  plates  IX 
and  XXXII  of  Science  Bulletin,  vol.  XL  At  that  time  no  ob- 
servations had  been  recorded  on  the  mating.  The  male  mounts  the 
female,  grasping  her  with  the  middle  pair  of  legs,  the  first  pair 
flexed  beneath  him,  and  the  abdomen  somewhat  to  the  left  side. 
This  decided  and  invariable  position  to  the  left  is  due,  of  course, 
to  the  asymmetrical  structure  of  the  male  genitalia.  The  fre- 
quency and  duration  of  copulation  of  various  pairs  under  obser- 
vation is  given  herewith. 


148 


The  University  Science  Bulletin. 


MATING  OBSERVATIONS  COVERING  25  DAYS 

Pair  1. 

Pair  2. 

Pair  3. 

Pair  4. 

Pai 

r5. 

Pair  6 

Date. 

Obs. 
mat- 
ing. 

Hrs. 
dura- 
tion. 

Obs. 
mat- 
ing. 

Hrs. 
dura- 
tion. 

Obs. 
mat- 
ing. 

Hrs. 
dura- 
tion. 

Obs. 
mat- 
ing. 

Hrs. 
dura- 
tion. 

Obs. 
mat- 
ing. 

Hrs. 
dura- 
tion. 

Obs. 
mat- 
ing. 

Hrs. 
dura- 
tion. 

May  15 

May  17 

X 

X 

X 

X2 

X 

X 

X 

X2 

X 

X 

X 

4  + 
6+ 

2-1- 
2 

iV2+ 

X 

X 

X2 

X 

X 

X 

4-^ 

2  + 

X 
X 

4-f 

X 
X 
X 

X2 

X2 

X2 

X 

X 

4+ 
4+,2 

X 
X 
X 
X 

4+ 
6+ 
1 

l}/2 

X 

4+ 

May  18 

May  19 

2+ 

2H+ 

5-t- 

X 

2+ 

X 
X 

X 
X 

X2 

1-1- 

May  20 

6+ 

May  21 

X 

4H+ 

2 

May  22 

May  24 

X 

1 

X 
X 

V4 

X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 

May  25 

May  27 

X 

X 

May  28 

X 
X 

X 

May  29 

- 

May  31 

X 
X2 
X 
X 

1-h 

2+ 

X 
X 
X 

June  1 

3M 

X 

X 

7+ 
2+ 

June  2 

X 
X 

4+ 

X 

2+ 

1M+ 

June  3 

June  4 

X 
X 

June  7 

X 

X 

June  9 

X 

In  the  table  X  signifie.s  observed  mating, 
plus  sign  indicates  that  the  bags  were  mating 


If  followed  by  the  figure  2  it 
when  first  or  last  observed,  or 


indicates  two  separate  matings.    The 
both. 


The  above  table  indicates  that  the  mating  clasp  may  last  as  long 
as  seven  hours,  that  matings  are  frequent,  often  twice  a  day,  and 
that  they  occur  almost  daily  over  a  considerable  period  of  time. 
Matings  were  observed  in  the  laboratory  from  May  to  November. 

OVIPOSITION. 

The  eggs  are  either  completely  or  partly  buried  in  the  sand,  ceph- 
alic end  uppermost.  The  partly  buried  dry  egg  is  white  and  very 
difficult  to  discover  in  dry  sand.  When  it  is  moistened  it  changes  to 
an  amber  or  ferruginous,  and  with  the  sand  takes  on  a  deeper  colored 
and  glistening  appearance,  which  further  adds  to  its  resemblance  to 
a  sand  grain.  A  photograph  of  the  egg  upon  a  background  of  sand 
is  shown  on  plate  XIII.  The  number  of  eggs  deposited  by  a  female 
during  the  season  must  be  200  or  more,  because  192  nymphs  and  eggs 
have  been  recorded  for  one  female,  and  it  is  quite  likely  that  I  did 


Hungerfokd:    Tiik  Toad  Bug.  149 

not  count  the  full  coniplenicnt.  Oviposition  continues  throughout 
the  season,  the  number  of  eggs  laid  per  day  varying  from  1  to  13, 
from  2  to  6  being  perhaps  the  average.  A  study  of  the  hatching 
dates  on  the  life-history  tables  will  indicate  the  rate  of  egg-laying 
very  well. 

INCUBATION. 

The  incubation  period  varied  from  twelve  to  fifteen  days.  The 
red  eyespots  of  the  embryo  are  evident  a  number  of  days  before 
hatching.  The  egg  increases  in  size  somewhat  as  the  embryo  devel- 
ops, and  the  egg  becomes  darker  with  development. 

HATCHING. 

I  was  fortunate  enough  to  observe  the  hatching  process  with  the 
binoculars  on  several  occasions,  but  never  in  as  satisfactory  lighting 
as  I  should  have  liked.  The  cephalic  end  of  the  egg  shell  splits  longi- 
tudinally, squarely  between  the  eyes,  and  extends  back  above  the 
dorsum  of  the  embryo.  Through  this  rent  a  white,  bulging  body 
appears,  resembling  the  bubble  found  above  the  head  of  Corixa. 
The  front  part  of  the  head  of  the  embryo  pulsates  rapidly.  By  slow 
straining  heaves  the  embryo  crowds  out  through  the  opening.  Its 
body  is  as  soft  and  pliable  as  a  caterpillar.  By  bulging  the  fore  part 
of  the  body  and  contracting  the  latter  part,  it  worms  its  w^ay  to  free- 
dom— a  creamy-white  creature  marked  wdth  two  large,  dark-red 
eyes,  and  v/ith  a  body  nearly  cylindrical  in  shape  and  the  thick 
limbs  all  most  economically  tucked  away  upon  its  venter.  Then 
comes  the  postnatal  molt,  the  casting  of  the  shroud  that  binds  the 
embryo.  Standing  erect  upon  its  caudal  end,  its  body  encompassed 
and  its  limbs  tied  down  by  a  diaphanous  membrane  that  still  holds 
it  helpless  to  the  empty  casque  in  which  it  was  formed,  it  struggles 
for  freedom.  First  the  membrane  gives  way  above  the  head  and 
emergence  begins.  As  this  skin  slips  back,  the  knob-like  antennse, 
which  were  directed  downward  along  the  beak,  are  free  and  change 
their  position.  Then  the  beak  appears,  and  after  slowly  bending 
back  and  forward,  one  front  leg  pops  out  free,  then  the  other.  Fi- 
nally all  the  legs  are  free,  and  the  little  bug  settles  down  upon  them, 
the  shroud  still  about  the  tip  of  the  abdomen.  He  flattens  out  into 
a  toad  bug,  and  after  resting  for  a  time,  as  if  the  birth  struggles  had 
been  most  exhausting,  he  suddenly  becomes  lively  and  starts  away. 
Over  the  first  moist  pebble  he  passes,  the  molt  is  left,  if  by  chance  it 
did  not  remain  fastened  in  the  slit  of  the  egg-shell.  A  period  of 
thirty  minutes  often  is  consumed  in  the  hatching.     One  bug  that 


150  The  University  Science  Bulletin. 

hatched  at  2:35  p.  m.  was  creamy  white  with  dark-red  eyes  at  3:12 
p.  m.,  when  a  faint  pattern  began  to  appear.  At  3:35  it  was  some- 
what darker  and  the  pattern  more  distinct.  Thus  it  takes  an  hour 
or  so  for  it  to  attain  its  characteristic  color. 

MOLTING. 

An  examination  of  the  life-history  tables  will  indicate  that  mor- 
tality at  molting  time  was  very  great  in  the  rearings.  This  suggests 
that  conditions  were  not  favorable  to  normal  molting.  In  the  light 
of  R.  Takahashi's  observations  on  the  Ochteridae,  published  in  Jap- 
anese,* it  may  be  that  Gelastocoris  nymphs,  like  those  of  Ochterus 
formosanus  Mats.,  normally  fashion  for  themselves  small  cells  of 
sand  above  the  ground  in  which  the  moltings  take  place.  The  sand 
in  the  rearing  jars  was  packed,  and  perhaps  too  coarse  for  the 
nymphs. 

The  nymph  becomes  very  plump  of  body  along  toward  the  time 
to  change.  Several  first-instar  bugs  were  under  observation  during 
the  process  of  molting.  The  bug  rests  upon  the  sand,  all  legs  out- 
spread and  apparently  rigid.  A  longitudinal  rent  appears  on  the 
dorsum  of  the  head  and  thorax,  and  the  greenish  or  creamy-white 
nymph  begins  bulging  out,  the  dorsal  part  of  the  thorax  leading. 
Shortly  the  head  is  freed,  the  slit  through  the  old  skin  extends  back 
to  the  abdomen  and  then  laterally  to  the  margins  of  the  body.  These 
lateral  fissures  enable  the  bug  to  work  his  way  out.  The  new  form 
is  so  much  larger  than  the  old  exuvium  which  encased  it,  that  one 
wonders  how  it  was  ever  tucked  away  in  so  small  a  fepace.  When 
the  new  instar  is  entirely  free,  the  old  skin  may  snap  back  in  place 
again  and  look  like  a  perfect  nymph,  save  that  the  eyes  are  whitish 
instead  of  dark  red. 

The  following  article  appeared  in  Japanese,  by  R.  Takahashi. 
The  observations  are  so  interesting,  and  relate  to  a  family  so  little 
known,  that  English-reading  students  will  be  glad  to  give  Mr.  Taka- 
hashi credit  for  it. 

These  observations  pertain  to  Ochterus  jormosanus  (Mats.),  wliich  is  not 
uncommon  in  Formosa. 

1.  The  adults  Hve  upon  the  sandy  shores  of  ponds  and  streams,  where  the 
colors  of  their  backs  merge  into  their  surroundings,  rendering  them  difficult  to 
discover.  They  are  not  able  to  submerge  and  do  not  run  out  upon  the  water, 
where  they  are  occasionally  found  by  accident;  but  the  nymphs  are  amphib- 
ious, being  often  seen  submerged. 

*  R.  Takahashi:  "Observations  on  the  Ochteridte,"  Trans.  Nat.  Hist.  Soc.  Formosa,  vol. 
XI,  No.   55,   pp.   119-125   (1921). 


Hungerford:    The  Toad  Bug.  151 

2.  The  adults  arc  very  active,  while  the  nymphs  are  rather  inactive.  The 
death  ft^igning  has  never  been  observed. 

3.  The  species  is  not  gregarious,  but  two  or  three  nymphs  are  sometimes 
found  in  groups. 

4.  The  n>-inphs  sometimes  \ibrate  the  abdomen  vertically  for  a  few  seconds 
when  resting  on  the  shore. 

5.  The  nymphs  cover  their  backs  completely  with  sandy  granules.  All  the 
nymphal  instars  have  this  habit.  Their  heads  are  provided  on  the  front  with 
12  to  14  stout  processes  jM'ojecting  forward  and  arranged  in  a  transverse  row, 
with  which  they  scoop  the  sand  upon  the  heads  and  push  them  backward  by 
the  front  legs. 

6.  The  m-mphs  construct  for  themselves  small  cells  of  sand  above  the 
groimd,  using  the  processes  on  the  front,  in  which  the  meltings  take  place. 
All  the  instars  have  this  peculiar  habit. 

7.  The  nymphs  with  the  dorsums  wettable  are  amphibious  in  habit,  being 
often  found  submerged.  When  submerged,  the  bodies  are  held  always  just 
below  the  surface  film,  and  they  swim  rather  awkwardly,  moving  all  the  legs, 
but  do  not  swim  deeper.  A  store  of  air  for  respiration  when  submerged,  is 
carried  with  the  insect  on  the  lower  surface  of  the  abdomen,  and  the  nymphs 
now  and  then  turn  on  their  backs  at  the  surface,  thus  exposing  the  lower  sur- 
face of  the  abdomen  into  free  air  to  take  new  supply  of  air.  This  is  done  very 
quickly. 

8.  The  mating  habit  is  almost  as  in  the  insects  of  Microvelia,  but  the 
males  do  not  remain  on  the  backs  of  their  mates  for  a  long  time  when  the 
copulation  is  finished.    The  males  and  females  mate  repeatedly. 

9.  The  eggs  are  singly  placed  upon  the  sandy  granules,  or  upon  the  decayed 
leaves  on  the  ground. 

10.  The  egg  is  similar  in  shape  to  that  of  Gelastocoris,  species  figured  by 
Doctor  Hungerford  (1919),  measuring  about  0.7  mm.  in  length. 

11.  There  are  five  nymphal  instars,  as  is  common  for  other  Heteroptera, 
and  the  nymphal  stage  lasts  more  than  one  month. 

12.  In  the  adults  the  front  and  middle  tarsi  are  two-jointed,  and  the  hind 
three-jointed,  while  in  the  nymphs  all  the  tarsi  are  always  two-jointed. 

13.  The  adults  may  be  seen  near  Taihoku  at  almost  any  time. 

NOTES  ON  REARINGS. 
FIRST  PAIR. 
Rearing  Number  4- 
This  pair  consisted  of  the  female  that  I  had  kept  in  the  laboratory- 
all  winter  and  a  male  selected  from  the  spring  shipment.     They 
were  confined  in  a  six-inch  stender  in  which  had  been  placed  a  layer 
of  sand  with  a  place  scooped  out  on  one  side  for  water.    The  water, 
however  was  taken  up  by  the  sand  and  the  entire  mass  became 
water-soaked.     To  make  a  dry  footing  for  tlie  bugs,  a  bit  of  cork 
was  set  upright  in  the  soil,  but  this  too  became  wet.     Green  algae 
covered  the  sand,  the  cork,  the  sides  of  the  jar,  and  even  the  bugs 
were  green  with  it.    The  surface  of  the  sand  was  found  on  July  1 


152  The  University  Science  Bulletin. 

to  be  teeming  with  nematodes  tliat  were  intent  upon  consuming 
the  fly  carcasses  supplied  as  food  for  the  bugs:    No  careful  examina- 
tion was  made  of  this  jar  until  July  1,  when  four  white  eggs  were 
found  on  top  of  the  cork,  placed  there  in  an  endeavor,  apparently, 
to  keep  them  out  of  the  wet,  soggy  sand.    I  then  supplied  dry  sand 
to  reduce  the  moisture,  and  on  July  6   found  eggs  in  tlie  sand; 
observed  mating  on  that  date  also.    July  12  two  nymphs  hatched. 
July  20,  there  were  several  active  first-instar  nymphs.     July  24, 
removed  18  dead  first-instar  forms  and  observed  nine  live  ones. 
There  were  several  eggs  yet  to  hatch  and  some  fresh  eggs.    July  27, 
removed  two  second-instar  forms  and  took  out  18  dead  first-instar 
bugs.    Seven  live  ones  were  observed  at  this  time.    July  30,  observed 
a  couple  hatch  and  counted  10  active  first-instar  nymphs;  also  re- 
moved 7  dead  ones.     On  August  1  there  were  nine  live  first-stage 
forms;  removed  2  dead  ones.    August  2,  counted  11  live  first-stage 
bugs  and  removed  6  dead  ones.    On  the  3d  removed  2  dead  ones.  On 
the  4th  there  were  3  newly  hatched  bugs,  and  I  removed  5  dead 
ones.    On  August  5  one  first-instar  bug  was  observed  feeding  upon 
another.     August  7  there  were  at  least  18  nymphs,  three  of  them 
white,  denoting  recent  hatching.     August  8,  7  dead  firsts  were  re- 
moved, but  a  dozen  were  still  lively.     August  9  another  freshly 
emerged  nymph  was  noted,  and  on  the  10th  2  more.     August  13, 
removed  a  second-instar  form,  and  on  August  15  another  newly 
hatched  was  noted.     August  16,  a  dark  nymph  was  caught  in  the 
act  of  killing  a  white  newly  emerged  brother.     He  was  upon  the 
back  of  his  victim  with  beak  inserted  just  back  of  the  unfortunate's 
head.     The  latter  was  struggling,  but  to  no  avail.     On  August  18 
one  second-instar  form  w'as  isolated  and  34  dead  first-stage  bugs 
removed.     Mr.  Hoffman  found  three  nymphs  feeding  upon  their 
fellows,  and  all  three  victims  were  alive  and  kicking.    There  were 
about  a  dozen  live  bugs  of  this  stage.     August  21'  brought  forth 
3  second-stage  bugs,  which  were  taken  out.     August  22,  11  dead 
firsts  were  moved,  and  on  the  23d  3  dead  firsts  and  1  live  second 
removed.    August  24,  2  dead  firsts  were  taken  out.     August  29,  5 
dead  first  were  removed,  and  on  August  31  the  female  was  found 
dead.     Twenty-three  dead  nymphs  were  counted  out  and  8  eggs 
transferred  to  another  stender.    No  live  nymphs  were  present,  and 
the  jar  was  set  aside  till  September  20,  when  a  careful  count  was 
made  of  remains — 10  eggs  and  16  nymphs  completed  the  record  for 
this  pair.    The  above  notes  have  been  given  to  show  the  result  of 
trying  to  rear  the  bugs  together. 


Hungerford:    The  Toad  Bug. 


153 


This  pair  of  bii^s  was  observed  mating  May  15,  17,  18,  19,  20,  21, 
22,  24,  27,  June  2,  7,  22,  and  July  6.  No  count  of  eggs  laid  was 
attempted,  but  22  eggs,  160  first-instar  and  10  second-instar  nymphs 
were  taken,  a  total  progeny  of  192,  with  egg-lay  noted  from  July  1 
to  last  of  August — this  the  performance  of  a  female  that  spent  the 
preceding  winter  in  the  laboratory. 


REARING 

RECORD  OF  NO. 

4. 

No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. 

1 

Days  in  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

4a 

July  12 

July  21 

Aug.    2 

Aug.  15 

Aug.  30 

Sept.  19 

& 

9 

12 

13 

15 

20 

69 

Oct.     6 

4b 

July  12 

July  20 

Died 
July   24 

8 

4° 

4c 

July  16 

July  24 

Aug.  13 

Died 
Aug.  16 

8 

20 

3° 

.... 

4d 

July   16 

July  25 

Aug.    5 

Aug.  16 

Aug.  31 

Sept.  23 

cf 

9 

11 

11 

15 

24 

70 

Nov.  17 

4e 

Aug.     1 

Aug.  13 

Died 
Aug.  31 

12 

18° 

4f 

Aug.    8 

Aug.  18 

Died 
Aug.  31 

10 

13° 

4g 

Aug.  20 

Sept.  10 

Died 
Sept.  14 

21 

4° 

4h 

Aug.  21 

Sept  11 

Sept.  20 

Oct.     5 

Died 
Oct.  20 

21 

9 

16 

15° 

71? 

4i 

Aug.  21 

Sept.   6 

Sept.  17 

Sept.  27 

Oct.  26 

& 

16 
1° 

11 

10 

29 

76? 

Oct.  28 

4j 

Aug.  23 

Died 
Sept.  24 

°  indicates  "died"  in  this  and  following  tables. 

SECOND  PAIR— OFFSPRING  IN  FOUR  SERIES. 
Rearing  Number  1,  Series  m. 
On  July  6,  Mrs.  Wiley  brought  me  7  bugs  from  Chanute,  which 
she  had  captured  July  3.  One  pair,  observed  mating,  I  placed  in  a 
small  stender  on  sand  with  a  sprig  of  moss.  July  8,  quite  a  number 
of  eggs  were  in  the  sand,  so  the  adults  were  removed  to  another 
stender.  On  July  15,  the  eggs  showed  pink  eyespots  of  the  embryo 
within;  and  on  July  19,  8  hatched  and  were  isolated  for  rearing. 
July  21,  5  more  hatched;  and  July  24,  3  more.  Thus  at  least  16 
eggs  were  laid  by  this  female  in  two  days,  and  the  incubation  period 
was  12  or  13  days.  The  rearing  table  marked  Im  series  is  a  record 
of  fifteen  of  these  nymphs. 


154  The  University  Science  Bulletin. 

Rearing  Number  1,  Series  n. 
The  pair  was  transferred  from  Im  to  this  stender,  containing 
sand,  on  July  8 ;  observed  mating  on  the  11th  and  removed  on  the 
12th.    Could  find  no  eggs,  and  though  this  stender  was  studied  till 
July  29,  no  nymphs  appeared. 

Rearing  Number  1,  Series  o. 
Transferred  the  pair  from  number  In  to  this  stender,  containing 
sand,  July  12.  On  July  24,  4  nymphs  hatched,  and  so  I  removed  the 
adults  to  another  stender  and  transferred  the  nymphs  to  jars  as  they 
hatched.  The  rearing  table  marked  lo  series  indicates  the  dates  of 
hatching.  But  to  this  list  of  49  must  be  added  the  following:  July 
27,  6  first  instars  in  large  stender,  labeled  lOh.  August  6,  4  dead 
instars;  and  August  8,  2  more.  A  total,  therefore,  of  61  hatched 
from  eggs  deposited  between  July  12  and  24,  an  average  of  5  per 
day,  with  a  record  of  13  for  August  6.    Incubation  period,  12  days. 

Pair  Number  Ip. 

This  pair  of  adults  was  transferred  from  lo  on  July  24.  They 
were  observed  mating  on  this  date  and  again  on  August  8,  when  2 
first-instar  forms  hatched.  Removed  the  male  on  August  10  and 
returned  on  the  18th,  This  female  was  caught  twice  feeding  upon 
her  own  offspring.  Besides  the  65  nymphs  hatched  from  August  8 
to  September  27  and  used  in  rearings,  28  dead  ones  were  taken  out 
of  this  stender  in  which  the  pair  was  confined — a  total  of  93  off- 
spring between  July  24  and  September  27.  The  observed  matings 
were  on  July  24,  August  8,  August  21  and  September  20.  One  adult 
died  October  4  and  the  other  October  23.  The  rearing  data  are  pre- 
sented on  that  portion  of  the  table  marked  IP  series. 

Summing  up  for  this  pair  of  bugs  taken  from  the  wild  on  July  3 
and  entered  for  observation  on  July  6,  we  get  a  total  progeny  of  170 
hatched,  and  of  this  number  we  were  able  to  rear  to  the  adult  stage 
4  insects. 


Hungerford:    The  Toad  Bug. 


155 


SECOND  PAIR,  REARING  TABLE. 

Series  Im. 


No. 

Date 

hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. 

Days  in  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

Inift 

July  19 

Died 
Aug.  14 

26° 

Imb 

July  19 

Died 
Aug.    7 

19° 

July  19 

Aug.  11 

Oct.     7 

Died 
Oct.  25 

23 

27 

19° 

Imd 

July  19 

Aug.    6 

Died 
Aug.  12 

18 

6° 

Ime 

July  19 

Died 
Aug.    5 

17° 

Imf 

July  19 

Died 
Aug.    9 

21° 

Img 

July  19 

Died 
July  28 

9° 

Imh 

July  21 

Aug.    1 

Aug.  11 

Died 
Aug.  16 

11 

10 

5° 

Ipii 

July  21 

July  28 

Aug.  11 

Died 

Aug.  16 

7 

14 

5° 

Imi 

July  21 

Aug.    7 

Died 
Aug.  11 

17 

4° 

Imk 

July  21 

Died 
Sept.  12 

53° 

1ml 

July  21 

Died 
July  30 

9° 

Imm 

July  21 

July  25 

Aug.    6 

Aug.  15 

Aug.  30 

Sept.  19 

cf 

4 

12 

9 

15 

20 

60 

Nov.   4 

Imn 

July  24 

Aug.    9 

Died 
Aug.  10 

16 

1° 

Imo 

July  24 

Died 
July  30 

6° 

i 

156 


The  University  Science  Bulletin. 


Series  lo. 


No. 

Date 

hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. 

Days  in  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

loa 

July  24 

Aug.  11 

Died 

Aug.  14 

18 

2.° 

lob 

July  24 

Aug.  10 

Died 
Aug.  13 

17 

3° 

July  24 

Died 
July  30 

6° 

lod 

July  24 

Died 
July  31 

7° 

July  25 

Died 
Aug.  10 

16° 

lof 

July  25 

.Aug.  12 

Sept.    2 

Sept.  17 

Died 
Oct.     1 

18 

21 

15 

13° 

log 

July  25 

Aug.  10 

Died 

Aug.  14 

16 

4° 

loi 

July  29 

Died 

Aug.    8 

10° 

loj 

July  29 

Died 
Aug.    5 

- 

7° 

lok 

July  29 

Aug.  13 

Aug.  31 

Sept.  13 

Sept.  25 

Oct.  21 

9 

15 

18 

13 

12 

26 

84 

Dec.  20 

lol 

July  29 

escaped 

lom 

July  29 

Died 
.Aug.  11 

13° 

July  29 

Died 

Aug.    5 

7° 

July  29 

Died 
Aug.  10 

12° 

lop 

July  30 

Died 
Aug.    7 

8° 

loq 

July  30 

Aug.  13 

Sept.  15 

Died 

Sept.  24 

14 

33 

9° 

lor 

July  30 

Died 
Aug.    1 

2° 

los 

July  30 

Aug.  11 

Aug.  28 

Sept.  12 

Sept.  24 

Died 
Oct.   22 

12 

17 

15 

12 

29 

85 

lot 

July  30 

Died 

Aug.  28 

29° 

July  30 

Died 

Aug.    8 

9° 

July  30 

Died 

Aug.    8 

9° 

lox 

July  30 

Died 

Aug.    9 

10° 

loy 

July  30 

Died 

Aug.    8 

9° 

loaa 

Aug.    1 

Died 
Aug.    9 

8° 

Hungerford:    The  Toad  Bug. 


157 


Series  lo. — Concluded. 


No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. - 

Days  in  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

lobb 

Aug.    1 

Aug.  18 

Died 
Sept.  17 

17 

30° 

locc 

Aug.    1 

Died 
Aug.    8 

7° 

lodd 

Aug.    2 

Died 
.\ug.  10 

8° 

Aug.  2 

Died 
Aug.    9 

7° 

loff 

Aug.  2 

Died 
Aug.  9 

7° 

logg 

Aug.  2 

Died 
Aug.    9 

7° 

lohh 

Aug.  2 

Died 

Aug.    9 

7° 

loii 

Aug.  2 

Died 
Aug.    9 

7° 

lojj 

Aug.  2 

Died 
Aug.    9 

7° 

lokk 

Aug.  4 

Died 
Aug.  10 

6° 

loll 

Aug.  4 

Died 
Aug.  14 

10° 

Aug.  4 

Died 
Aug.    9 

5° 

Aug.  4 

Died 
Aug.    9 

5° 

Aug.  5 

Died 
Aug.  10 

5° 

lopp 

Aug.  5 

Died 
Aug.  15 

10° 

loqq 

Aug.  5 

Sept.  12 

Sept.  26 

Oct.  21 

Died 
Nov.  17 

38 

14 

25 

27° 

Aug.  6 

Died 
Aug.  11 

5° 

Aug.  6 

Died 
Aug.  10 

4° 

lott 

Aug.  6 

Died 
Aug.  13 

7° 

Aug.  6 

Died 
Aug.    9 

3° 

Aug.  6 

Died 
Aug.  12 

6° 

Aug.  6 

Died 
Aug.  1-1 

8' 

Aug.  6 

Died 
Aug.  IC 

4" 

. 

Aug.  6 

Died 
Aug.  IC 

7' 

. 

lobbb 

Aug.  6 

Died 
Aug.  1 

. 

5' 

158 


The  University  Science  Bulletin, 


Series  Ip. 


No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. 

Days  in  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

Ipa 

Aug.    8 

Died 
Aug.  14 

6° 

Ipb 

Aug.    8 

Died 
Aug.  14 

6° 

^ 

Idc 

Aug.    9 

Died 
Aug.  14 

5° 

Ipd 

Aug.    9 

Died 
Aug.  15 

6° 

Aug.    9 

Sept.  22 

Died 
Oct.   15 

44 

23° 

lof 

Aug.  10 

Died 
Aug.  18 

8° 

iDff 

Aug.  10 

Died 
Aug.  16 

6° 

Iph 

Aug.  10 

Died 
Aug.  19 

9° 

Ipi 

Aug.  10 

Died 
Aug.  19 

9° 

Ipj 

Aug.  10 

Died 
Aug.  17 

7° 

Ipk 

Aug.  11 

Died 
Aug.  12 

1° 

Ipl 

Aug.  12 

Died 
Aug.  25 

13° 

1pm 

Aug.  16 

Died 
Aug.  22 

6° 

Ipn 

Aug.  16 

Died 

Aug.  22 

6° 

8° 

Ipo 

Aug.  16 

Died 
Aug.  24 

• 

Ipp 

Aug.  16 

Died 
Sept.   9 

24° 

Ipq 

Aug.  16 

Died 
Aug.  23 

7° 

Ipr 

Aug.  16 

Died 
Sept.    2 

17° 

Ips 

Aug.  17 

Died 
Aug.  26 

9° 

Ipt 

Aug.  17 

Died 
Aug.  29 

12° 

Ipv 

Aug.  18 

Sept.  13 

Died 
Oct.     6 

26 

23° 

Ipw 

Aug.  19 

Died 

Aug.    1 

13° 

Ipx 

Aug.  21 

Died 

Aug.    2 

12° 

• 

ipy 

Aug.  21 

Sept.    9 

Sept.  23 

Oct.   11 

Died 
Oct.   23 

19 

14 

18 

12° 

Hungerford:    The  Toad  Bio. 


159 


Series  Ip — Continued. 


No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex 

Days  in  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

Ipz 

Aug.  22 

Died 
Aug.  11 

20° 

Ipaa 

Aug.  23 

Died 

Aug.    4 

12  = 

Ipbb 

Aug.  25 

Died 
.\ug.  16 

22° 

Ipcc 

Aug.  26 

Died 
Aug.  29 

3° 

Ipdd 

Aug.  27 

Died 
Sept.   4 

8° 

Ipee 

Aug.  28 

Sept.  13 

Died 
Sept.  24 

16 

11° 

Ipff 

Aug.  29 

Sept.  15 

Sept.  25 

Oct.   14 

Nov.  17 

Died 

Nov.  26 

17 

10 

20 

34 

9° 

Iphh 

Aug.  30 

Died 
Sept.   9 

10° 

loii 

Sept.   3 

Died 
Sept.  17 

14° 

Ipjj 

Sept.    4 

Died 

Sept.  24 

20° 

Ipkk 

Sept.   4 

Died 
Sept.  15 

11° 

IpU 

Sept.    7 

Died 
Sept.   9 

2° 



Innun 

Sept.    7 

Died 
Sept  15 

8° 

. 

Ipnn 

Sept.    9 

Sept.  21 

Oct.   15 

Nov.    4 

Died 
Dec.    4 

12 

24 

20 

30° 



Ipoo 

Sept.    9 

Died 

Sept.  18 

9° 

Ippp 

Sept     9 

Died 
Sept.  17 

8° 

Ipqq 

Sept.   9 

Died 
Sept.  19 

10° 

Iprr 

Sept.  10 

Died 
Sept.  21 

11° 

Ipss 

Sept.  12 

Died 

Sept.  22 

10° 

Iptt 

Sept.  12 

Died 
Sept.  26 

14° 

Ipuu 

Sept.  12 

Died 
Sept.  21 

9° 

Ipw 

Sept.  12 

Died 
Sept.  28 

16° 

Ipww 

Sept.  12 

Oct.     8 

Died 
Oct.   12 

26 

4° 

Ipxx 

Sept.  12 

Sept.  23 

Oct.    11 

Died 

Nov  13 

11 

18 

33° 

160 


The  University  Science  Bulletin. 


Series  Ip — Concluded. 


No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 

molt. 

4th 
molt. 

5th 
molt. 

Sex. 

Days  in  stages. 

Adult 

1st. 
10° 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

Ipvy 

Sept.  13 

Died 
Sept.  23 

Ipzz 

Sept.  13 

Died 
Sept.  21 

8° 

Ipaaa 

Sept.  13 

Sept.  29 

Died 
Nov.    2 

16 

3° 

Ipbbb 

Sept.  13 

Died 
Sept.  28 

15° 

Ipcee 

Sept.  15 

Oct.     5 

Nov.    2 

Died 
Dec.  30 

20 

28 

59° 

Ipddd 

Sept.  15 

Died 

Sept.  25 

10° 

- 

Ipeee 

Sept.  15 

Sept.  27 

Died 
Oct.   15 

12 

18° 

IpiTf 

Sept.  17 

Died 
Sept.  30 

13° 

lD2Ee 

Sept.  17 

Died 

Sept.  25 

8° 

■• 

Iphhh 

Sept.  17 

Died 
Sept.  19 

2° 

Ipiii 

S?pt.  22 

Died 
Oct.     2 

10° 

Ipjji 

Sept.  22 

Died 

Sept.  2S 

6° 

Ipkkk 

Sept.  23 

Died 
? 

? 

IpUl 

Sept.  23 

Died 
Oct.   13 

20° 

Ipmmm 

Sept.  23 

Oct.    18 

Died 
Oct.   30 

25 

12° 

Ipnim 

Sept.  25 

Dir-d 
Sept.  29 

4° 
4° 

Ipooo 

Sept.  27 

Died 
Oct.     1 

It  was  not  planned  when  I  began  the  studies  to  carry  the  rearings 
to  such  an  extent  and  therefore  the  simple  system  of  designating 
became  cumbersome. 


Hungerford:    The  Toad  Bug. 


161 


THIRD  PAIR. 


On  Alay  14  I  placed  a  pair  in  a  small  stendcr  on  the  sand.  Eggs 
showed  eyespots  June  2.  June  10,  4  nymphs  appeared,  and  on  this 
date  adult  female  died.  The  rearings  from  this  pair  are  given  in  the 
following  table: 


No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. 

Days  i  n  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

lb 

June  10 

June  28 

Died 
July  17 

18 

19° 

Ic 

June  10 

June  19 

July     1 

Escaped 
July     6 

9 
15° 

12 

■- 

Id 

June  13 

Died 
June  28 

le 

June  13 

Died 
June  21 

8° 

If 

June  14 

June  24 

July     3 

July  11 

July  19 

Aug.    2 

9 

10 

9 

8 

8 

12 

47 

Jan.  11, 
1921 

Ih 

June  17 

June  28 

Died 
June  28 

11° 

li 

June  17 

July     1 

Died 
July     6 

14 

5° 

Ij 

June  21 

Died 

July     2 

11° 

Ik 

June  21 

June  28 

July     6 

July  15 

July  22 

Aug.    7 

9 

7 

8 

9 

7 

16 

47 

Nov.    2 

11 

June  22 

July     5 

July  15 

July  23 

Aug.    6 

Aug.  31 

_9_ 

13 

10 

8 

14 

25 

70 

Oct.   10 

162 


The  University  Science  Bulletin. 


FOURTH  PAIR. 

In  a  large  stender  containing  sandy  loam,  and  labeled  No.  7,  was 
placed  a  mating  pair  of  bugs,  June  28.  Eggs  were  discovered  in  the 
soil  on  July  8.  July  13  nymphs  hatched  and  the  adults  were  re- 
moved. July  20,  7  more  nymphs  were  out.  These  12  nymphs  were 
isolated  in  stenders  and  a  tabular  report  of  these  follows.  The  in- 
cubation period  was  somewhere  between  7  and  15  days. 


No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. 

Days  in  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

7a 

July  13 

Escaped 

7b 

July  13 

July  20 

Died 
July  23 

7 

3° 
16 

8° 

7c 

July  14 

July  21 

Aug.    6 

Died 

Aug.  14 

7 
9° 

7e 

July  14 

Died 
July  23 

7f 

July  20 

Died 
July  30 

10° 

7g 

July  20 

July  28 

Aug.    6 

Aug.  16 

Sept.    1 

Sept.  21 

9 

8 

9 

10 

16 

20 

63 

Oct.  28 

7h 

July  20 

Aug.  16 

Aug.  30 

Sept.  12 

Sept.  22 

Oct.   17 

cf 

27 

14 

13 

10 

25 

89 

Nov.  14 

7i 

July  20 

Aug.    4 

Aug.  14 

Aug.  26 

Sept.  11 

Sept.  29 

& 

15 

10 

12 

16 

18 

71 

Nov.  21 

7j 

July  20 

Aug.    9 

Died 
Aug.  12 

20 

3° 

7k 

July  20 

Died 
July  28 

8° 

71 

July  31 

Aug.    6 

Aug.  16 

Sept.  14 

Died 
Sept.  27 

6 

n 

29 

13° 

Hungerford:    The  Toad  Bug. 


1H:} 


FIFTH  SERIES. 

A  sprig  of  moss  and  eartli  containing  eggs  were  plaeed  in  a  slender 
July  12.  Tiiese  eggs  were  deposited  by  bugs  sent  rae  July  6,  amongst 
the  material  in  which  they  were  packed.  They  began  hatching  July 
20  and  finished  July  24.    The  record  of  the  eight  nymphs  isolated  is 

sliown  in  tabular  form  below. 


No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. 

Days  in  stages. 

Adult 

1st. 

2i 

3d. 

4th. 

5th. 

Tot. 

died. 

9a 

July  20 

July  .30 

Aug.  19 

Sept.  IP 

Oct.     7 

Died 
Oct.   18 

10 

20 

27 

22 

11 

9b 

July  20 

Aug.    6 

Aug.  18 

Aug.  30 

Sept.  13 

Oct.     5 

& 

17 

12 

12 

14 

22 

77 

Oct.    17 

9c 

July  21 

Aug.    2 

Died 
Aug.  10 

12 

8 

9d 

July  21 

Died 
July  31 

10 

9e 

July  24 

Died 
Aug.    9 

16 

9f 

July  24 

Died 
Aug.    3 

10 

9g 

July  24 

Aug.    1 

Died 
Aug.    9 

8 

8 

.... 

9h 

July  24 

July  30 

Aug.  15 

Sept.    S 

Sept.  22 

Died 
Oct.   14 

6 

16 

25 

13 

22 

164 


The  University  Science  Bulletin. 


SIXTH  PAIR. 

Placed  2  female  bugs  in  large  stender  marked  No.  12,  on  July  20. 
These  bugs  were  from  those  Mrs.  Wiley  collected  July  3.  On 
August  2,  2  nymphs  appeared — an  incubation  period  of  13  days. 
The  nymphs  continued  to  appear,  and  their  record  is  shown  in 
table  12  below.  September  5  only  1  adult  was  to  be  found,  and 
yet  it  was  impossible  for  the  bugs  to  escape  from  the  jar.  Septem- 
ber 6  both  were  gone  from  view,  but  later  they  were  again  above 
the  sand.  In  cloudy,  threatening  weather,  the  toad  bugs  bury 
themselves  beneath  the  surface  and  appear  again  in  sunny  weather. 

Table  12. 


No. 

Date 
hatched. 

1st 
molt. 

2d 
molt. 

3d 
molt. 

4th 
molt. 

5th 
molt. 

Sex. 

Days  in  stages. 

Adult 

1st. 

2d. 

3d. 

4th. 

5th. 

Tot. 

died. 

12a 

Aug.    2 

Died 
Aug.    8 

6 

12b 

Aug.    2 

Died 
Aug.    9 

7 

12c 

Aug.    8 

Died 

Aug.  14 

6 

12d 

Aug.    9 

Died 

Aug.  14 

5 

12e 

Aug.  10 

Died 
Aug.  12 

2 

12f 

Aug.  10 

Aug.  26 

Died 
Sept.  17 

16 

22 

12g 

Aug.  11 

Died 
Aug.  19 

8 

12h 

Aug.  15 

Died 

Aug.  21 

6 

12i 

Aug.  15 

Died 

\ug   ?2 

7 

12j 

Aug.  16 

Died 

Aug.  28 

12 

SEVENTH  PAIR. 

In  stender  marked  2a,  placed  a  mating  pair  on  May  25.  On  June 
3,  found  an  egg  imbedded  in  the  sand  for  about  one-half  of  its 
length.  On  June  13,  found  11  eggs,  5  or  6  of  which  showed  pink  eye- 
spots,  so  removed  the  adults  to  another  stender.  The  eggs  began 
hatching  June  17  and  continued  until  June  28.  The  minimum  in- 
cubation period  for  some  of  the  eggs  was  15  days. 


Hungerkord:    The  Toad  Bug.  165 

EIGHTH  PAIR. 

Pair  from  Chanute  placed  in  stcnder  labeled  la,  June  2.  The  fe- 
male died  June  10.  Eggs  hatched  June  24  and  June  28.  Incuba- 
tion period  at  least  14  days,  likely  longer. 

NINTH  PAIR. 

Female  emerged  from  one  of  the  rearings  (11)  on  August  31,  from 
a  nymph  that  hatched  June  22.  Added  a  male,  but  no  mating  ob- 
ser\ed,  and  female  died  October  10. 

TENTH  PAIR. 

Female  emerged  from  one  of  the  rearings  (Ik)  on  August  10,  from 
nymph  that  hatched  June  21.  Placed  a  male  with  her  and  matings 
were  observed  on  August  13,  15  and  18.  No  eggs  w^ere  found  after 
most  thorough  search.    Female  died  November  4. 

DESCRIPTION  OF  STAGES. 

(See  plates  XIII  and  XIV.) 

THE  EGG. 

Size.    Length,  1.25  mm.;  diameter,  .91  mm. 

Shape.  Broadh'  oval.  The  sui-face  is  roughly  granular  and  marked  into 
regular  hexagonal  areas  bj'  thickened  ridges  of  the  chorion. 

FIRST-INSTAR  NYMPH. 

Size.    Length,  2  mm.;  width  of  thorax,  1.4  mm.;  width  of  head,  1.2  mm. 

Color.  Eyes  dark;  body  mottled  to  checkered  obscurely,  general  color 
sometimes  light,  sometimes  dark.  Legs  and  abdomen  are  usually  marked  a^ 
described  for  later  instars. 

Structural  peculiarities.  Eyes  larger  relatively  than  in  succeeding  instars. 
They  are  not  placed  upon  as  high  elevations  and  the  inner  emarginations  are 
not  marked.  The  beak  is  four-segmented;  the  antenuEe  three-segmented;  and 
the  tarsi  appear  one-segmented,  terminating  in  two  claws.  The  middle  and 
hind  tarsi,  however,  have  a  short  basal  segment,  making  two.  Lateral  margin 
of  mesothorax  less  in  length  than  that  of  metathorax. 

SECOND-INSTAR  NYMPH. 

Size.    Length.  2.5  mm.;  width  of  thorax,  1.9  mm.;  width  of  head,  1.5  mm. 

Color.  Eyes  still  dark,  not  distinctly  banded.  Pattern  obscured,  that  of 
limbs  and  abdomen  as  in  later  instars. 

Structural  peculiarities.  The  inner  emargination  of  the  eyes  a  little  more 
marked  than  in  the  preceding  instar.  Lateral  margin  of  mesothorax  equal  in 
length  to  that  of  metathorax. 


166  The  University  Science  Bulletin. 

THIRD-INSTAR  NYMPH. 

Size.     Length,  3.5  mm.;  width  of  thorax,  2.7  mm.;  width  of  head,  2.0  mm. 

Color.  The  eyes  faintly  banded  with  three  bars  of  color,  one  of  them  on 
the  inner  margins.  The  general  pattern  variable;  but  when  defined,  like 
other  instars. 

Structural  ])eculiarities.  The  inner  emargination  of  the  eye  now  regular 
and  marked.  Ocelli  faintly  visible  in  some,  while  in  others  unmistakably 
present.  Lateral  margin  of  Mesothorax  now  a  little  longer  than  that  of 
metathorax. 

FOURTH  INSTAR  NYMPH. 

Size.     Length,  4.5  mm.;  width  of  thorax.  3i.6  mm.,  width  of  head,  2.4  mm. 

Color.  The  general  pattern  as  in  fifth.  The  eyes  banded  by  four  faint  bars, 
one  of  which  is  on  inner  margin. 

Sttructural  'peculiarities.  The  lateral  margin  of  mesothorax  now  is  about 
twice  as  long  as  that  of  the  metathoracic  margin,  due,  of  course,  to  the 
lengthening  of  the  wing  pads.  The  body  is  covered  with  closely  set  short, 
stout,  appressed  spines,  which  show  more  plainly  than  in  the  preceding  instar. 
The  ocelli  show  plainly  now.  They  are  located  just  above  the  lateral  arms 
of  what  I  take  to  be  the  epicranial  suture.  The  nymphal  exuvium  always 
shows  a  Y-shaped  rent  on  the  head,  and  it  is  on  the  ui^per  margin  of  the 
lateral  arms  of  this  fissure  that  the  ocelli  are  located. 

FIFTH  INSTAR  NYMPH. 

Size.     Length,  6.2  mm.;  width  of  thorax,  5.0  mm.;  width  of  head,  3.0  mm. 

Color.  Color  variable  from  very  light  to  very  dark,  and  the  pattern  varying 
from  obscure  to  distinct.  The  eyes  have  five  dark  bars.  There  are  two  pairs 
of  black  dots  on  the  face,  one  above  the  other,  and  laterad  and  below  the 
ocelli.  A  median  black  dot  on  vertex.  The  pronotum  has  front  two-thirds 
of  lateral  margins  of  prothorax  darker.  There  is  a  pair  of  black  spots  caudo- 
laterally  of  each  pronotal  elevation,  of  which  there  are  two.  The  mesonotum 
has  two  rectangular  black  spots  on  front  margin  either  side  of  median  line. 
There  is  a  transverse  row  of  four  black  dots  across  caudal  third  of  mesonotum; 
wing  pad  has  a  dark  rectangular  spot  on  shoulder;  the  middle  of  the  pad 
crossed  by  another  dark  area;  tip  and  inner  margin  of  pad  dark.  Two  faint 
dark  irregular  spots  in  the  outer  third  of  wing  pad  gives  a  mottled  effect  to 
the  whole.  A  white  spot  is  found  on  second  abdominal  segment  either  side 
of  median  line.  A  row  of  dark  marginal  spots,  roughly  triangular,  cover  the 
front  two-thirds  of  the  margin  of  each  abdominal  segment.  There  are  also 
two  rows  of  dark  submarginal  spots,  faint  and  ill  defined.  The  legs  are  banded 
with  dark  bands.  The  hind  tibise  have  three  dark  bands  besides  being  dark 
at  the  ends. 

Structural  -peculiarities.  The  wing  pads  of  the  mesothorax  now  extend 
almost  to  caudal  margin  of  thorax,  nearly  obscuring  lateral  margin  of  meta- 
thorax. 


Hungerford:   The  Toad  Bug.  167 


SUMMARY. 

The  toad  bug  places  its  eggs  in  the  sand.  The  eggs  hatch  in  about 
12  days.  There  are  five  nymphal  instars.  Forty-nine  first-instar 
bugs  transformed  in  an  average  of  15  days  (the  shortest  time  4 
days  and  the  longest  44  days).  Thirty-three  second-instar  bugs 
transformed  in  an  average  of  16  days  (shortest  time  8  days  and  long- 
est 33  days).  Twenty-two  third-instar  bugs  averaged  15  days  (the 
shortest  time  8  days  and  the  longest  period,  not  counted  in  the 
average  because  it  failed  to  molt,  was  59  days).  Eighteen  fourth- 
instar  forms  averaged  15^/^  days  (minimum  7  days  and  maximum 
34  days).  Thirteen  fifth-instar  forms  averaged  22  days  (minimum 
12  and  maximum  29  days).  The  average  number  of  days  for  the 
thirteen  adults  to  develop  from  the  hatching  to  emergence  was  701/2 
days.  The  shortest  period  was  47  days  and  the  longest  89  days. 
By  adding  the  12  days  incubation  of  the  egg,  we  get  a  total  de- 
velopment period  of  from  60  to  100  days.  The  adult  female  may 
deposit  a  dozen  eggs  a  day,  but  would  average  perhaps  only  2  to  6 
over  the  period  of  two  or  more  months.  One  hundred  and  ninety- 
two  eggs  and  nymphs  were  counted  from  one  female  from  July  6 
to  September  27.  The  nymphs  possess  ocelli,  plainly  distinguished 
as  early  as  the  third  instar.  All  the  stages  are  predaceous  and  much 
like  their  parents  in  other  habits  as  well. 


168  The  University  Science  Bulletin. 


PLATE  XIII. 

Gelastocoru  oculatus  Fabr. 

Fig.  1.  Adult  bug  (not  oculatus,  but  an  undescribed  species  in  western 
Kansas)  upon  sand,  showing  how  its  mottled  pattern  makes  it  difficult  to 
discern. 

Fig.  2.  Ventral  view  of  above  species  of  bug.  Enlarged  photograph  show- 
ing the  bug  clasping  three  lace  bugs. 

Fig.  3.  First-instar  nymph,  Gelastocoris  oculatus  Fabr.  Enlarged  dorsal 
view,  same  species  showTi  in  figures  1  and  2. 

Fig.  4.    Another  toad  bug.    Enlarged  ventral  view,  showing  bug  with  prey. 

Fig.  5.    First-instar  nymi)h,  Gelastocorifi  oculat^is  Fabr.,  ventral  view. 

Fig.  6"  Microphotograj)!!  of  egg  and  first-instar  nym])h.  Nymph  in  center 
of  picture  and  the  white  oval  egg  to  the  right  of  it.  The  egg  resembles  the 
sand  grains  very  closely. 


PLATK  XIII. 


(169) 


PLATE  XIV. 

Gelastocoris  ociilatii.'<  Fabr. 

Fig.  1.  Egg. 

Fig.  2.  First-instar  nymph. 

Fig.  3.  Second-instar  nymph. 

Fig.  4.  Third-instar  nymph. 

Fig.  5.  Fourtli-instar  nymph. 

Fig.  6.  Fifth-instar  nymph. 

(170) 


.?■• 


Hungerford:    The  Toad  Bi(;. 


171 


PLATE  XIV. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  6— October,  1922. 

(Whole  Series,  Vol.   XXIV,  No.   6.) 

ENTOMOLOGY  NUMBER  V 


CONTENTS: 

A  New  Subterranean  Isopod  (Crustacea)...//.  B.  Hungerford. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  po.st   office  in  Lawrence  as  second-cla.ss  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

^■oL.  XIV.J  October,  1922.  [No.  6. 


A  New  Subterranean  Isopod  from  Kansas. 

Ccecidotea   tridentata    (Crustacea). 

By  H.  B.  HUNGERFORD, 

Professor  of  Entomology,  University  of  Kansas. 

IN  MARCH  of  1919,  Mr.  William  Hoffmann,  field  assistant  in  our 
department  of  entomology  at  the  University  of  Kansas,  brought 
to  me  for  determination  some  specimens  of  an  isopod  which  he 
had  taken  from  a  cistern  in  Lawrence,  Kan. 

They  prove  to  belong  to  a  new  species  of  the  genus  Coecidotea. 
For  them  I  propose  the  name  Coecidotea  tridentata,  because  the 
propodus  of  the  first  pair  of  legs  of  the  male  is  armed  with  three 
conspicuous  processes,  a  character  which  separates  them  from  the 
previously  described  species. 

The  crustacean  genus  Coecidotea  Pack.,  as  the  name  implies,  is 
characterized  by  the  absence  of  eyes,  by  the  fact  that  the  terminal 
segment  of  the  body  is  much  longer  than  broad,  and  by  the  elongate, 
narrow  body.  An  analytical  key  to  the  genus  was  given  by  Harriet 
Richardson  in  her  monograph  of  the  isopods  of  North  America,  in 
1905.  There  were  known  at  that  time  four  species,  namely,  C.  stygia 
Pack.,  C.  nickajackensis  Pack.,  C.  richardsonce  Hay  and  C.  smithsii 
Ulrich.  Doctor  Ortmann,  1918,  in  chapter  XXV  of  "Fresh-water 
Biology,"  had  Miss  Richardson's  work  in  mind  when  he  said  there 
were  four  species  of  the  genus  and  that  they  are  found  in  caves, 
springs  issuing  from  caves,  and  artesian  wells.  However,  in  1911, 
in  the  Pomona  College  Journal  of  Entomology,  volume  3,  No.  3, 
Blanche  Stafford  described  a  fifth  species,  namely,  Coecidotea  ala- 
bamensis,  from  a  well  in  Auburn,  Ala. 

The  following  table  will  serve  to  separate  the  six  species  of  the 
genus  now  known: 

(175) 


176  The  University  Science  Btlletin. 

A.    Propodus  of  first  pair  of  legs  armed  witli  one  or  more  triangular  processes. 

B.  Propodus  of  first  pair  of  legs  armed  with  a  triangular  process  near 
the  distal  end  and  with  a  long  spine  at  the  proximal  extremity. 
Uropods  about  one-half  the  length  of  terminal  abdominal  segment. 
Outer  branch  three-fourths  as  long  as  inner,  which  equals  the 
peduncle  in  length.  C.  nickajackennis  Pack. 

BB.    Propodus  of  first  pair  of  legs  armed  with  two  triangular  processes. 

C.  Propodus  with  three  additional  short  processes.  Uropods 
about  as  long  as  terminal  abdominal  segment.  Outer  branch 
two-thirds  as  long  as  inner,  which  is  two-thirds  as  long  as 
jieduncle.  C.  stygia  Pack. 

CC.    Propodus  with  three  additional  spines  not  jjrocesses.  Uropods 

a    little    longer    than    terminal    abdominal    segment.      Outer 

branch  about  one-half  as  long  as  inner,  which  is  two-thirds 

as  long  as  peduncle.  C.  alabamohsis  Stafford. 

BBB.    Projjodus  of  first  jjair  of  legs  armed  with  three  triangular  processes. 

C.  tridentata,  sp.  nov. 

AA.    Propodus  of  first  pair  of  legs  not  armed  with  triangular  processes,  but 
edged  inside  with  spines. 

B.  First  pair  of  antenna\  with  flagellum  composed  of  eleven  articles, 
extend  one-third  the  length  of  the  fifth  article  of  the  peduncle 
of  the  second  antenna.  Second  antenna  longer  than  the  body; 
flagellum  composed  of  about  eighty-six  articles. 

C.  richardsonce  Hay. 
BB.    Fii-st   pair  of  antennae,  with  flagellum   composed   of  five  articles, 
extend   half  the   length   of  the   peduncle   of   the   .second  antenna. 
Second  pair  of  antennae  "probably  as  long  as  body,"  flagellum  com- 
posed of  "at  least  forty  segments."  C .  smithsii  Ulrich. 

Coecidotea  tridentata  sp.  nov. 

(Plate  XV.) 

Size.  The  bod.\'  without  the  antennae  and  uiopoda  measures  in  length  from 
9  mm.  to  19  mm.  and  in  width  from  1%  mm.  to  3  mm.  The  length  of  the 
body  is  approximately  five  or  s-'x  t  ines  the  width.  This  species  is  much  larger 
than  the  others  of  this  genus  that  have  been  describsd.  From  the  descriptions 
I  infer  that  10  mm.  is  about  the  maximum  of  (*.  styi/ia  Packard,  the  largest 
member  of  the  genus,  while  3  mm.  is  supposed  to  be  the  maximum  of  C. 
smithsii  Ulrich,  the  smallest.  These  figures  suggest  that  the  smallest  mature 
C.  triden'ata  are  about  the  size  of  some  of  the  members  of  other  species,  but 
the  laigpst  indi\iduals  are  fully  double  that  size. 

Color.  The  color  is  chalky  white,  the  body  wall  being  sufficiently  clear 
to  show  the  dark  food  canal  within.  Material  .stored  in  alcohol  appears  very 
[lale  yellowish  gray. 

Stmchire.  Head:  Narrower  than  first  thoracic  segment.  Wider  than  long. 
The  cephalic  margin  narrower  than  the  caudal,  somewhat  concave,  and  bear- 
ing the  antennules  and  antennae,  the  bases  of  the  latter  appearing  veiy  heavy 
when  compared  with  the  size  of  the  head.  The  antennule  consists  of  the 
basal  segments  and  a  flagellum  of  from  twelve  to  eighteen  segments,  the  two 
parts  of  about  equal  length;  the  basal  segment  stoutest,  a  trifle  longer  than 
twice  its  width;  second  segment  two-thirds  as  broad  as  basal  and  about 
.same  length;  third  segment  much  smaller;  five-eighths  as  broad  and  one- 
half  as  long  as  second.     Its  distal  end  bears  the  tai)ering  flagellum.     The  an- 


Hungerford:   A  New  Isopod.  177 

remui'  arc  r(>lati\  ely  lar^r  and  consi.-^t  of  a  ba.^al  part  of  six  segments  and  a 
flagelhim  of  from  sixty  to  eifihty  segments.  Each  of  the  first  four  segments 
of  the  basal  ]iart  is  broader  than  h)ng.  Taken  together  they  are  equal  in 
length  to  th(>  fifth  segmtmt.  wiiicii  is  a  little  shorter  than  the  sixth,  from  which 
arises  the  many-segmented  flagellum.  The  mandible  bears  a  large  three-seg- 
mented flattened  jialp  and  two  chitin-tipped  processes,  one  a  chisel-hke  cutting 
edge  and  the  other  bearing  from  four  to  seven  teeth. 

Thorax:  The  segments  of  the  thorax  are  loosely  articulated  and  their 
lateral  margins  are  fringed  with  ver^•  short,  stout  setae.  All  are  broader  than 
long.    It  bears  seven  pairs  of  legs,  of  which  the  first  pair  is  subchelate. 

The  first  pair  of  legs  is  shorter  than  the  others.  In  the  males  the  propodus 
is  very  large  and  bears  three  well-develoi)ed  processes,  one  at  the  base  and 
two  near  the  distal  end.  The  ba^sal  one  is  bifurcate  in  some  and  in  others 
bears  instead  a  strong  seta.  There  are  seven  divisions  to  each  limb,  counting 
the  clawlike  terminal  one.  The  propodus  is  the  enlarged  fifth  division  by  this 
count.  The  limbs  bear  many  strong  setse  and  increase  in  length  from  the 
first  to  the  last. 

Abdomen:  The  first  two  segments  of  the  abdomen  are  short.  The  so-called 
third  is  nearl\-  twice  as  long  as  wide  and  carries  the  uropods,  each  of  wiuch 
consists  of  a  basal  part  and  two  terminal  branches.  The  uropods  are  longer 
than  the  abdominal  segment  which  bears  them,  the  relative  length  being  5:3. 
The  basal  segment  is  nearly  as  long  as  the  last  abdominal  segment,  the  ratio 
about  6:7  in  the  males.  The  two  branches  are  of  very  unequal  length;  the 
one  female  possessing  inopods  had  this  basal  segment  1:3,  the  inner  being 
much  the  larger.  The  relative  lengths  vary  from  3:2  in  the  female  to  4:1  in  the 
male.  The  inner  branch  is  to  the  peduncle  as  3:4.  There  is  considerable  varia- 
tion in  the  comparative  lengths  of  these  parts.  The  second  pleopod  of  the 
male,  the  first  of  the  female  and  the  third  pleojiods  of  both  sexes  are  unlike 
those  figured  by  StafYord  for  C.  alnbamensi.<<. 

Holotype,  allotype  and  jiaratypes  in  alcohol.    Kansas  University'  collection. 

Tiie  females  are  smaller  than  the  males  and  do  not  have  as  well- 
developad  propodi.  The  sexual  dimorphism  appears  not  to  have 
been  recorded  in  the  genus  heretofore.  Another  point  not  mentioned 
in  descriptions  is  the  fact  that  the  females  possess  the  flattened 
brood  pouches  or  oostegites  at  the  base  of  the  first  four  pairs  of  legs. 
Our  specimens  were  obtained  in  June,  and  some  of  the  females  bear 
these  plates. 

I  asked  Mr.  Hoffmann,  who  gathered  the  material,  to  submit  a  few 
notes  relating  the  circumstances  of  their  collection.  His  notes  in 
substance  follow: 

The  cistern  frcni  which  these  specimens  were  taken  is  about  eight 
feet  in  diameter  and  nine  feet  deep.  It  contains  a  square  brick 
filter,  resting  on  the  bottom  in  the  center,  which  measures  three  or 
four  feet  square  at  base  and  tapers  to  its  top,  some  four  feet  above, 
where  it  is  just  large  enougli  for  a  fotn'-inch  casing,  which  extends 


178  The  University  Science  Bulletin. 

above  to  within  three  feet  of  the  top.  This  casing  surrounds  the 
pipe  leading  to  the  pump,  and  is  open  at  the  top.  The  water  supply- 
ing the  cistern  is  caught  upon  the  roof  of  the  house  and  conducted 
to  the  cistern  by  galvanized  pipes  three  or  four  inches  in  diameter. 

On  one  occasion  during  a  rain  two  isopods  were  observed  by  the 
lady  of  the  house  to  be  washed  out  of  the  elbow  pipe  leading  from 
the  gutter  along  the  eaves  of  the  house  onto  the  sloping  tin  roof  of 
the  kitchen,  thence  into  another  gutter  and  down  the  pipe  to  the 
cistern.  She  concluded,  therefore,  that  these  animals,  which  she 
pumped  up  by  way  of  the  pitcher  pump  in  the  kitchen  sink,  were 
either  "rained  or  had  bred  in  the  collection  of  wet  leaves  in  the  gut- 
ters of  the  house  or  in  the  elbow  of  the  pipe  leading  from  them." 

A  number  of  specimens  were  taken  alive  in  the  water  pumped 
from  the  cistern.  One  of  these  was  placed  in  a  specimen  jar,  three 
and  one-half  inches  in  diameter  and  three  inches  deep,  where  it 
lived  in  one  and  one-half  inches  of  water  from  June  18  until  July  26. 
The  water  was  replenished  from  time  to  time  with  dirty  pond  water, 
containing  many  small  organisms. 

Most  of  the  specimens  died  within  a  few  days.  When  several 
were  placed  together  they  seemed  to  take  no  notice  of  each  other. 
The  pleopods  were  observed  to  be  in  vibration  as  an  individual 
made  its  way  through  the  water. 

It  is  unfortunate  that  we  were  too  busy  to  run  any  behavior  exper- 
iments upon  these  most  interesting  forms. 

Note. — Through  the  kindness  of  the  custodian  of  Crustacea  I  had  the 
privilege  of  examining  the  Coecidotea  material  in  the  National  Museum  at 
Washington,  D.  C.  One  jar  marked  Coecidotea  stygia  contains  eight  vials; 
four  of  these  contain  large  specimens  which  belong  to  the  species  I  have  de- 
scribed as  new.  It  is  interesting  to  note  that  they  were  collected  at  Topeka, 
Kan.  They  bear  labels  as  follows :  "Gift  of  E.  A.  Popenoe,  Topeka,  Kansas,"  and 
were  taken  "April  9,  May  4,  May  12,  May  29,  1912."  The  other  four  vials  con- 
tain material  taken  from  "Graham's  Spring,  Lexington,  Va.,  1876";  Richard- 
son's Spring,  Ky.,  W.  P.  Hay,  Col.'';  "Irvington,  Ind.,  from  wells  W.  P. 
Hay";  "Mammoth  Cave,  Ky.,  R.  E.  Call." 

The  last  four  lots  are  much  smaller  specimens  than  the  Kansas  material. 
The  material  from  Virginia,  Richardson's  Spring,  Ky.,  and  from  Indiana,  differs 
materially  from  the  Kansas  species.  The  species  is  broader  than  the  Kansas 
crustacean,  and  the  third  pleopods  are  not  only  much  broader  comparatively, 
but  are  more  truncate  at  the  tip. 


1(S0  TriK    UXIVERSITV    SCIEN(,'K    BULLETIN. 


PLATE  XV. 

Ccecidotea  tridentata    (Crustacea). 

Fig.  1.    First  pleopod  of  female. 

Fig.  2.    First  antenna. 

Fig.  3.    Mandible. 

Fig.  4.    First  pair  pleopods  of  male. 

Fig.  5.   Leg  of  female  bearing  oostegite. 

Fig.  6.    Second  pair  pleopods  of  male. 

Fig.  7.    Front  leg  of  male. 

Fig.  8.   Adult  male. 

Fig.  9.    Third  pleopod  of  male. 

Fig.  10.    Front  leg  of  female. 

Fig.  11.    Mature  female  carrying  brood  pouch. 


HrN(;KHFOHD:    A  New  Tsopod. 


181 


PLATE  XV. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  7— October,  1922. 

(Whole  Series,  Vol.  XXI^^  No.  7.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

Studies  on  Cicadella  Hieroglyphica  (Homoptera), 

Lucy  M.  Hackman. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


1/ 


TABLE  OF  CONTENTS. 

PAGE 

Introduction 189 

Life  history  notes 190 

Description  of  the  species 190 

Hosts 192 

Hibernation 192 

Spring  appearance 192 

Mating  and  oviposition 192 

Nymphs , 193 

Description  of  instars 194 

Adults 195 

Morphological  studies 195 

General  morphology  of  the  abdomen 195 

Development  of  the  male  genitalia 196 

Development  of  the  female  genitalia: 198 

(187) 


I 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  7. 


Studies  on  CicadeUa  hierogjyphica  Say  (Homoptera, 

Cicadellidae.) 

BY  LUCY  M.  HACKMAN. 

Submitted  to  the  department  of  entomology  and  to  the  graduate  faculty  of  the  University 
of  Kansas  in  partial  fulfillment  of  the  requirements  for  the  degree  of  master  of  arts,  May  IS, 
1922. 

INTRODUCTION. 

THE  following  notes  on  the  life  history  of  CicadeUa  hieroglyphica 
consist  of  observations  made  from  specimens  in  the  field  and  in 
the  laboratory.  A  growth  of  yomig  willows  along  the  Kansas  river 
offered  a  splendid  opportunity  for  the  former,  for  there  CicadeUa 
hieroglyphica  may  be  found  at  all  seasons  in  very  large  numbers. 
The  laboratory  observations  were  made  from  specimens  collected  at 
this  place  and  reared  on  willow  in  the  laboratory.  The  most  satis- 
factory results  M^ere  obtained  when  the  willow  was  planted  in  large 
glass  rearing  cages.  The  leaf  hoppers  could  move  about  at  will  and 
were  easily  observed. 

Only  a  general  description  of  the  abdomen  is  given,  for  the  chief 
concern  of  this  paper  is  the  genitalia.  In  tracing  out  the  develop- 
ment of  the  genitalia  in  the  male  and  female,  the  adult  genitalia 
were  used  as  a  starting  point.  To  trace  the  development,  the  various 
changes  in  the  genitalia  from  one  instar  to  another  were  studied.  A 
study  of  the  ventral  surface  of  the  eighth  and  ninth  abdominal  seg- 
ments was  sufficient  in  the  case  of  the  female,  for  all  three  genital 
appendages  are  readily  seen  from  a  ventral  view.  But  in  the  case 
of  the  male,  where  the  two  pairs  of  appendages  are  dorsal  in  position 
only,  the  development  of  the  ventral  ones  could  be  traced  by  such  a 
study.  Therefore,  particular  attention  was  given  to  these  dorsal  or 
internal  genitalia.     For  this  purpose  the  pygofers  were  split  open 

(189) 


190  The  University  Science  Bulletin. 

along  their  dorsal  surface,  the  overlying  integument  carefully  re- 
moved, and  the  genitalia  thus  exposed.  Great  modifications  in  the 
genitalia  occur  within  a  single  instar,  and  an  attempt  was  made  to 
give  a  rather  detailed  account  of  these  modifications  in  the  fifth 
instar. 

The  writer  wishes  to  express  her  appreciation  to  all  who  have  as- 
sisted her  in  this  work.  Professor  Hunter  has  always  been  most 
kind  in  helping  in  whatever  way  possible.  Dr.  Paul  B.  Lawson, 
under  whose  direction  the  work  was  done,  has  given  freely  of  his 
time  and  experience.  Kathleen  Doering,  Philip  Readio  and  Robert 
Guntert  are  also  deserving  of  thanks  for  their  interest  and  assist- 
ance. 

LIFE  HISTORY  NOTES. 

The  following  are  some  of  the  references  to  this  species: 

Tettigonia  hieroglyphica  Say,  Jl.  Acad.  Nat.  Sci.  Phila.,  vi,  p.  313,  1831. 

Tettigonia  hieroglyphica  Sig*>  Ann.  Soc.  Ent.  Fr.,  ser.  3,  iii,  p.  805,  1855. 

Tettigonia  hieroglyphica  G.  and  B.,  Hemip.  Colo.,  p.  81,  1895. 

Tettigonia  hieroglyphica  Ball,  Proc.  la.  Acad.  Sci.,  viii,  p.  51,  1901. 

Tettigoniella  hieroglyphica  Van  D.,  Trans.  San  Diego  Soc.  Nat.  Hist.,  ii, 
p.  52,  1914. 

Tettigoniella  hieroglyphica  Be  L.,  Tenn.  St.  Bd.  Ent.,  Bui.  17,  p.  20,  1916. 

Cicadella  hieroglyphica  Van  D.,  Cat.  Hemip.  N.  A.,  p.  597,  1917. 

Cicadella  hieroglyphica  01s.,  Bui.  Am.  Mus.  Nat.  Hist.,  xxxvii,  p.  3,  1918. 

Cicadella  hieroglyphica  Lawson,  Kan.  Univ.  Sci.  Bui.,  xii,  p.  85,  1920. 

DESCRIPTION  OF  THE  SPECIES. 

The  following  is  the  original  description: 

Tettigonia  hieroglyphica.  Dull  rufous;  head  and  scutel  lineated;  heme- 
lytra  spotted. 

Inhabits  Arkansas. 

Body  obscurely  dull  rufous;  head  with  a  black  dot  at  tii),  above  literate 
with  black;  thorax  with  a  dusky  jiosterior  disk;  scutel  with  black  more  or  less 
curved  lines;  hemelytra  obsoletelj'  spotted,  nervures  being  pale;  beneath  pale 
yellowish;  pectus  with  large  black  spots;  feet  immaculate;  tergum  blue-black, 
edge  yellow.    Length  to  tij)  of  hemelytra  one-fifth  of  an  inch. 

Dr.  P.  B.  Lawson,  in  his  paper  on  the  Cicadellidse  of  Kansas,  gives 
the  following  description  of  the  species: 

Form.  Rather  stout.  Length,  6  to  7  mm.  Vertex  bluntly  conical,  wider 
than  long.  Pronotum  nearly  twice  as  wide  as  long,  posterior  angles  broadly 
rounded,  posterior  margin  medially  emarginated.  Elytra  broad,  but  exceeding 
the  abdomen. 

Coloi^.  Varying  from  brick  red  to  greenish  and  slaty  blue.  Black  markings 
on  vertex  very  strong  and  distinct,  enclosing  a  light-colored  T  on  basal  half. 
Elytra  with  pale  bands  along  the  costal,  claval  and  sutural  margins. 


HaCKMAN  :     ClCADELLA  HiEROGLYPHICA.  ,  191 

External  genitalia.  Fcmalo:  Last  ^•cntl•al  sefinient  about  as  wide  as  long, 
lateral  margins  triangularly  luoduccd;  pygofers  long  and  narrow,  equaling  or 
slightlj'  exceeding  ovipositor,  bearing  a  few  stout  hairs.  Male:  Last  ventral 
segment  less  than  twice  as  wide  as  long;  plates  long,  broad  at  base,  but 
tapering  to  long  acute  apices,  margins  fringed  with  short  hairs;  jiygofers  long 
and  narrow,  equaling  or  exceeding  plates  and  bearing  stout  hairs. 

Internal  male  genitalia.  Styles  short,  distinctly  bent  in  at  point  of  attach- 
ment to  connective  by  a  large,  heavily  chitinized  lobe,  then  curving  outward 
and  tapering  gradually  to  bhmt  apex,  with  an  outwardly  projecting  process; 
connective  slender,  Y-shaped,  stem  of  Y  broadening  to  broad  base;  oedagus 
with  pair  of  short  processes  extending  dorsad  from  its  point  of  attachment  to 
connective,  a  long  process  leaving  it  dorsally  from  a  point  a  little  past  its 
middle,  and  a  similar  longer  one  leaving  it  apically,  the  latter  to  the  left  of 
the  former.  These  two  processes  are  narrow  and  long,  narrowest  at  the  base, 
and  widening  to  a  point  shortly  before  the  apex,  where  they  are  the  widest, 
the  right  one  wider  than  the  left  one,  and  then  tapering  to  the  acute  tips.  A 
pair  of  somewhat  narrow  triangular  chitinous  processes  extend  from  the  base 
of  the  anal  tube  to  the  main  body  of  the  oedagus. 

//o.s"/.^.    Taken  abundantly  on  willows. 

The  following  variety  occurs  along  with  the  typical  forms: 

Cicadella  hieroglyphica  var.  dolobrata  (Ball).     Its  bibliography 

follows : 

Tettigonia  hieroglyphica  var.  dolobrata  Ball,  Proc.  la.  Acad.  Sci.,  p.  52,  pi. 
3.  fig.  2,  190L 

Tettigonia  hieroglyphica  var.  dolobrata  DeL.,  Tenn.  St.  Bd.  Ent.,  Bui.  17,  p. 
20,  1916. 

Cicadella  hieroglyphica  var.  dolobrata  Van  D.,  Cat.  Hemip.  N.  A.,  p.  597, 
1917. 

Cicadella  hieroglyphica  var.  dolobrata  01s.,  Bui.  Am.  Mus.  Nat.  Hist., 
xxxviii,  p.  3,  1918. 

Cicadella  hieroglyphica  var.  dolobrata  Lawson,  Kan.  Univ.  Sci.  Bui.,  xii, 
p.  86,  1920. 

Doctor  Lawson  describes  this  variety  as  follows: 

This  is  a  smaller  form  than  the  preceding,  appearing  more  robust.  In  color 
it  is  typically  black,  retaining  a  few  of  the  lighter  markings  of  the  typical 
hieroglyphica  on  the  front,  vertex,  pronotum  and  scutellum,  and  generally 
having  the  claval  sutures  light. 

Genitalia  as  in  the  preceding  form. 

Distribution:    Occurs  along  with  the  typical  form. 

Hosts:     Willows. 

DISTRIBUTION. 

Doctor  Ball  gives  the  following:  "This  species,  as  a  whole,  is 
very  variable  in  size  and  color  and  recalls  Oncometopia  undata  and 
lateralis  in  their  red,  green  and  black  forms.  The  varieties  readily 
fall  into  two  series  on  structm-al  characters.  The  first  has  hiero- 
glyphica  and  dolobrata  as  the  extreme  in  darkening  up.     These 


192  The  University  Science  Bulletin. 

forms  are  the  only  ones  found  in  the  Mississippi  valley  and  as  far 
west  as  central  Kansas;  they  occur  also  in  Texas,  Arizona,  and 
Mexico." 

Van  Duzee  reports  it  from  Kansas,  New  Mexico,  Texas,  Illinois^ 
Missouri,  Iowa,  Nebraska,  and  Arizona. 

HOSTS. 

Cicadella  hieroglyphica  may  be  found  on  several  hosts.  Willows 
(Salix  longi folia  and  Salix  amygdaloides)  are  the  most  common  of 
these,  but  it  is  frequently  found  on  poplar  {Populus  monilifera). 
Occasionally  it  has  been  taken  on  the  broad-leafed  milkweed 
{Asclepias  syriaca)  and  on  giant  ragweed  (Ajnbrosia  trifida) . 

HIBERNATION. 

During  the  winter  the  adults  hide  among  the  fallen  leaves  and 
rubbish  on  the  ground,  and  appear  very  sluggish  when  disturbed. 
On  mild,  sunshiny  days  in  January  numbers  have  been  observed 
sunning  themselves  upon  the  stems. 

SPRING  APPEARANCE. 

About  the  middle  of  February,  or  when  the  willows  are  first  be- 
ginning to  bud,  the  greater  number  are  to  be  found  on  the  branches 
and  stems  of  the  willows.  They  are  very  gregarious,  and  often  are 
so  clustered  together  as  to  completely  hide  the  stem.  At  this  time 
of  year  they  feed  by  sucking  the  sap  from  the  stems,  and  give  off 
honeydew  in  such  quantities  as  to  cause  a  noticeable  spray.  Upon 
close  observation  this  honeydew  is  seen  to  be  given  off  in  a  rapid 
succession  of  droplets  from  the  anal  tube.  Several  specimens  were 
timed  in  the  operation,  and  from  fifteen  to  thirty  drops  were  given 
off  per  minute.  This  continues  for  several  hours  at  a  time  while  the 
insect  is  feeding.  Frequently  this  operation  is  accompanied  by  a 
spasmodic  raising  and  lowering  of  the  wings,  movements  of  the 
abdomen,  and  stroking  of  the  wings  and  abdomen  by  the  long  met- 
athoracic  legs. 

MATING  AND  OVIPOSITION. 

(Plate  XVII.  figs.   1-4.) 

Early  in  April  mating  takes  place.  By  this  time  the  willow  buds 
are  beginning  to  unfold  and  oviposition  begins.  Numbers  have  been 
watched  ovipositing,  both  in  the  field  and  in  the  laboratory,  and  the 
following  observations  made. 

The  eggs  are  inserted  in  tlie  tissues  of  the  upper  surface  of  the 
leaf  just  under  the  epidermis.     In  the  act  of  ovipositing  the  female 


Hackmax:    Cicadella  Hi?:roglyphica.  193 

braces  herself  firmly,  at  times  using  her  beak  in  addition  to  her 
legs  for  this  purpose.  In  all  eases  observed  she  always  worked  head 
up.  She  first  unsheaths  her  ovipositor,  punctures  the  epidermis  with 
its  tip,  and  then  inserts  it  to  its  full  length.  The  flat  surface  of  the 
ovipositor  now  rests  parallel  to  the  flat  surface  of  the  leaf,  with  its 
toothed  edge  pointing  forward.  Sawing  the  ovipositor  back  and 
forth  she  increases  the  size  of  the  slit  until  it  is  large  enough  for  the 
egg.  The  egg  then  passes  between  the  valves  of  the  ovipositor  into 
the  chamber  prepared  for  it,  and  the  ovipositor  is  withdrawn  and 
sheathed.    In  a  very  few  seconds  the  process  is  begun  again. 

The  time  taken  up  in  preparing  the  chamber  and  depositing  the 
egg  varied  in  several  cases  observed  from  two  to  five  minutes,  most 
of  which  time  was  spent  in  preparing  the  chamber.  To  cite  a  char- 
acteristic case,  the  whole  operation  occupied  two  and  one-half 
minutes,  the  two  minutes  being  spent  in  preparing  the  chamber  and 
the  half  minute  in  placing  the  egg. 

The  eggs  may  be  laid  singly  or  side  by  side  in  even  rows.  The 
largest  number  found  in  a  single  row  was  twenty-five  and  the 
largest  number  in  a  single  leaf  was  thirty-five.  The  eggs  in  a  hun- 
dred rows  or  masses  were  counted,  and  the  average  number  per  row 
was  found  to  be  seven. 

The  effect  of  oviposition  on  the  leaves  is  noticeable.  The  greater 
number  of  eggs  are  laid  in  leaves  not  fully  developed,  and  the  pres- 
ence of  the  eggs  causes  the  growing  leaf  to  become  distorted  and  to 
curl  around  the  eggs.  However,  in  no  case  observed  did  oviposition 
kill  the  leaf.     Eggs  have  been  found  in  both  willow  and  poplar 

leaves. 

NYMPHS. 

(Plate  XVI,  figs.   1-6.) 

Soon  after  oviposition,  nymphs  may  be  found  feeding  on  the 
leaves.  Eggs  observed  in  the  laboratory  hatched  in  from  eight  days 
to  two  weeks.  During  their  nymphal  life  these  little  leaf  hoppers 
molt  five  times  and  become  adult  in  a  little  over  two  months,  or  about 
the  middle  of  June.  By  the  middle  of  May  the  adults  of  the  over- 
wintering generation  are  all  dead.  Shortly  after  becoming  adult,  the 
new  adults  mate  and  another  generation  is  completed  by  the  end  of 
summer.  This  generation  consists  of  the  overwintering  individuals. 
Xymphs  af  this  generation  have  been  found  abundantly  on  giant 
ragweed  and  goldenrod,  and  in  all  probability  the  eggs  for  this  gen- 
eration are  laid  in  these  hosts. 


194  The  University  Science  Bulletin. 

DESCRIPTION  OF  THE  INSTARS. 

For  the  description  of  the  variout^  instars  an  attempt  was  made  to 
select  an  average  indivdual.  It  is  possible  to  separate  the  males  and 
females  of  the  same  instar  by  an  examination  of  the  ventral  surface 
of  the  eighth  and  ninth  abdominal  segments.  This  is  discussed  later 
in  greater  detail  under  the  development  of  the  genitalia  of  each  sex. 
Except  for  this,  and  a  slight  difference  in  size,  the  female  being  the 
larger,  the  two  sexes  are  practically  the  same. 

Egg. 

(Plate  XVII,  fig.  1.) 

Length,  1.25  mm.  Greatest  width,  .41.  Subovoid  in  shape,  somewhat  pointed 
at  one  end;  greenish  yellow  at  first;  just  before  hatching  deeper  yellow  with 
dark  e3'espots. 

First  Instar. 

(Plate  XVI,  fig.   1.) 

Length,  2.65  mm.  Width  across  eyes,  .65  mm.  Pale  yellowish  white,  eyes 
black  and  prominent;  anterior  margin  of  head  evenly  rounded;  two  pairs  of 
wing  pads  present,  first  pair  short,  barely  covering  base  of  second  pair,  caudal 
margin  in  form  of  an  inverted  W,  with  median  projection  extending  farther 
caudad  than  lateral  projections,  exposed  portion  of  second  pair  more  than 
twice  as  long  as  first  pair,  caudal  margin  nearly  straight,  the  segments  marked 
off  by  light  brownish  bands. 

Second  Instar. 

(Plate  XVI,  fig.  2.) 

Length,  3.15  mm.  Width  across  eyes,  .9  mm.  Color  same  as  in  first  instar; 
margin  of  head  similar;  wing  pads  have  increased  in  length  and  width,  but 
relative  shape  and  position  are  the  same;  caudal  margin  of  second  pair  bent 
slightly  cephalad  medially. 

Third  Instar. 

(Plate  XVI,  fig.  3.) 

Length,  4.25  mm.  Width  across  eyes,  1  mm.  Color  and  shape  of  head  un- 
changed; wing  pads  show  decided  change  in  length  and  width,  lateral  angles  of 
first  pair  produced  caudad  and  much  longer  than  median  projection;  exposed 
length  of  second  pair  but  little  longer  than  first  pair. 

Fourth  Instar. 

(Plate  XVI,  fig.  4.) 

Length,  5.3  mm.  Width  across  eyes,  1.35  mm.  Color  and  shape  of  head 
practically  the  same  as  in  preceding  instars;  lateral  angles  of  first  pair  of  wing 
pads  now  reach  almost  to  the  apex  of  the  second  pair;  lateral  angles  of  second 
pair  have  now  extended  farther  caudad,  making  median  indentation  in  caudal 
margin  more  pronoimced;  do  not  extend  beyond  second  segment  of  abdomen. 


Hackmax:    Cicadella  Hiekoglyphica.  195 

Fifth  Instar. 

(Plate  XVI,  fig.  .■).■) 
Lciititli,  0.4  nun.  Width  across  eyes.  1.4  nun.  Color  the  r^amc,  head  becom- 
ing!: slifrhtly  more  pointed,  markings  mon-  distinct  ;  lateral  angles  of  first  pair  of 
wing  iiads  still  further  jiroduced  and  are  now  about  the  same  length  as  second 
pair;  lateral  angles  of  second  pair  also  further  produced  caudad  and  now  extend 
almost  to  fourth  abdominal  segment. 

Adults. 

(Plate  XVI,  fig.  0.) 

The  seconci  generation,  or  the  overwintering  one,  consists  of  the 
typical  reddish  adults.  Th?  sexes  can  be  easily  distinguished  one 
from  the  other  by  the  darker  color  of  the  male  abdomen  as  con- 
trasted with  the  lighter  color  of  the  female's.  These  produce  the 
slaty-gray  individuals  of  the  summer  generation.  The  black  form, 
the  variety  dolobrata,  is  also  present  at  this  season.  However,  only 
males  of  this  form  have  been  found.  They  mate  with  the  slaty-gray 
individuals,  which  are  for  the  most  part  females,  although  males  of 
this  type  are  numerous.  Very  evidently,  the  dark  forms  are  di- 
morphic males  of  the  summer  generation.  A  single  dark  male  was 
taken  April  1  with  the  overwintering  red  forms,  but  whether  this  is 
one  of  the  summer  forms  which  has  overwintered  or  one  of  the  win- 
tering generation  which  differs  in  color  from  the  rest  of  the  gener- 
ation has  not  been  determined. 

MORPHOLOGICAL  STUDIES. 
GENERAL  DESCRIPTION  OF  THE  ABDOMEN. 

(Plate  XVIII,  figs.  1-3;    (5-9.) 

The  abdomen  is  joined  broadly  to  the  metathorax.  It  continues 
at  the  same  width  for  about  two-thirds  of  its  length,  and  from  there 
tapers  to  a  somewhat  pointed  apex.  In  a  general  cross  section  it  is 
semicircular  in  outline.  The  sternite  and  the  pleurites,  ventral  in 
position,  form  the  straight  part  of  the  semicircle,  while  the  curved, 
dorsal  tergite  forms  the  circular  part.  Eleven  segments  can  be  ac- 
counted for.  In  the  male,  six  of  these  are  represented  by  complete 
sternites.  pleurites  and  tergites,  but  in  the  female  only  five  are  so 
represented.  In  the  first  two  segments  only  the  sterna  and  terga  are 
present,  the  pleura  being  represented  by  pleural  membranes.  Seg- 
ments three  to  eight  in  tiie  male  and  three  to  seven  in  the  female  are 
typical.  The  terminal  segments  in  both  sexes  are  modified.  These 
modifications  are  discussed  under  the  heading  "external  genitalia" 
in  the  description  of  the  species.  In  addition  to  what  is  given  there, 
I  sliould  like  to  add  that  the  ventral  valve  of  the  adult  is  present, 
but  concealed  by  the  last  ventral  segment. 


196  The  University  Science  Bulletin. 

DEVELOPMENT  OF  MALE  GENITALIA. 

(Plate  XIX,  figs.  2,  4,  6,  8,  10,  12;   Plate  XX,  figs.   1-18.) 

The  male  genitalia,  consisting  of  three  pairs  of  valves  arise  from 
a  genital  area  on  the  ninth  abdominal  segment.  In  the  first,  second 
and  third  instars  there  are  two  pairs  of  valves  present.  These  de- 
velop in  small  chitinous  pockets,  which  are  attached  at  the  caudal 
edge  of  the  genital  area  with  their  apices  directed  caudad.  The 
genital  area  increases  slightly  in  size  in  each  successive  instar,  as  do 
the  pockets.  The  pockets  are  placed  one  upon  the  other,  the  ven- 
trally  placed  pocket  producing  the  plates  of  the  adult  and  the  more 
dorsal  pair  the  oedagus.  The  ventral  pair  is  about  twice  as  long  as 
the  dorsal.  Both  taper  caudad,  and  are  somewhat  rounded  at  the 
apex.  Each  pair  is  divided  into  its  right  and  left  valve  by  a  chit- 
inous median  partition. 

In  the  fourth  instar  there  is,  as  in  the  former  instars,  a  noticeable 
increase  in  size,  both  of  the  genital  area  and  of  the  ventral  and  dor- 
sal pockets.  In  addition  to  these  structures,  there  is  now  present  a 
pair  of  small  lateral  pockets  located  at  the  lateral  margin  of  the 
genital  area,  and  extending  caudad  as  far  as  the  bases  of  the  dorsal 
pockets.  In  these  lateral  pockets  develop  the  styles  of  the  adult 
genitalia.  In  the  fifth  instar  there  is  no  great  change  in  the  relati^'e 
position  and  shape  of  the  pockets.  There  is,  of  course,  a  natural 
increase  in  the  size  of  all  three  pockets. 

In  the  fifth  instar  the  greatest  changes  in-  the  developing  genital 
appendages  take  place.  The  three  pairs  of  valves  may  be  traced 
through  several  distinct  phases  of  development  by  a  study  of  the 
soft,  white  integument  which  can  be  drawn  from  the  chitinous  pock- 
ets. Details  of  five  particular  phases  might  be  mentioned.  (Plate 
XX,  figs.  6-18.) 

The  ventral  plates  which  develop  in  the  ventral  pockets  show 
very  little  change  throughout  the  successive  phases  of  this  instar. 

The  styles  which  develop  in  the  lateral  pockets  are  present  in  all 
five  phases  and  show  a  gradual  increase  in  size  through  the  phases. 
In  the  fourth  phase  they  first  show  their  permanent  attachment  to 
the  ventral  plates,  which  is  more  apparent  in  the  fifth  phase  and  in 
the  adult.  They  are  attached  on  their  outer  edges  near  the  base  of 
the  plate. 

The  developing  oedagus  or  the  integument  drawn  from  the  dorsril 
pockets  shows  the  most  remarkable  alterations.  In  phase  1  the 
cedagus  consists  of  two  valves  placed  parallel  to  the  ventral  plates. 


Hackman:    Cicadella  Hieroglyphica.  197 

In  phase  1,  ventral  aspect,  the  valves  placed  side  by  side  are  com- 
paratively narrow  at  their  bases,  widest  at  a  point  a  little  before  the 
middle,  and  then  tapered  to  somewhat  pointed  apices.  Dorsally 
they  do  not  appear  to  extend  as  far  cephalad  as  they  do  ventrally, 
nor  are  thej^  divided  into  right  and  left  valves  except  for  a  short 
distance  apically,  at  which  point  they  are  widely  separated. 

In  phase  2,  viewed  ventrally,  the  two  valves  have  become  longer 
and  slightly  narrower  than  in  the  preceding  phase,  and  instead  of 
their  former  relationship,  side  by  side,  the  right  valve  at  the  base  is 
now  assmning  a  ventral  position  with  respect  to  the  left  valve.  Also, 
the  flat  surfaces  of  the  valves,  instead  of  being  parallel  to  the  ventral 
plates,  are  now  assuming  a  perpendicular  position.  Between  the 
valves,  at  a  point  a  little  beyond  the  middle,  a  short,  slender  finger- 
like process  is  visible.  Viewed  dorsally,  this  process  appears  be- 
tween the  two  valves  at  the  point  of  wide  separation  in  phase  1  and 
projects  for  a  short  distance  caudad.  It  is  an  evagination  of  the 
integument  of  the  valves. 

In  phase  3,  ventral  aspect,  the  two  valves  have  continued  to  in- 
crease in  length.  The  right  valve  is  distinctly  folded  over  the  left 
valve  at  the  base.  A  lateral  aspect  shows  plainly  the  relative  posi- 
tion of  the  median  process  and  the  valves.  In  phase  3,  dorsal  aspect, 
there  is  little  change  in  the  valves  except  an  increase  in  length.  The 
median  process  also  shows  an  increase  in  length  and  width. 

In  phase  4,  ventral  view,  the  valves  again  are  longer  than  in  the 
preceding  phase,  and  the  median  process  also  shows  a  distinct  in- 
crease in  length.  The  folding  of  the  right  valve  over  the  left  valve 
is  more  complete,  and  the  bases  of  the  two  valves  are  farther  apart 
than  in  the  preceding  phase,  the  left  valve  having  moved  caudad. 

In  phase  5  the  oedagus  appears  very  similar  to  the  adult  oedagus. 
Here  the  valves  have  become  very  much  longer  than  in  phase  4. 
The  median  process  has  increased  greatly  in  length. 

During  the  development  of  the  valves  their  apices  have  main- 
tained their  relative  length  in  respect  to  the  ventral  plates.  The  in- 
crease in  length  has  been  taken  up  by  increased  dorsal  curvature. 

For  a  description  of  the  adult  male  internal  genitalia,  reference 
may  be  made  to  the  description  of  the  species. 


198  The  University  Science  Bulletin. 

DEVELOPMENT  OF  THE  FEMALE  GENITALIA. 

Ovipositor  of  Adult. 

(Plate  XVIII.  figs.  4-6.) 

In  the  female  the  genitaha  also  consist  of  three  pairs  of  append- 
ages. They  are  the  ventral  and  dorsal  pairs,  which  make  up  the 
ovipositor  proper,  and  the  lateral  pair,  within  which  the  ovipositor 
is  folded.  The  ventral  pair  arise  from  the  eighth  sternum,  and  the 
dorsal  and  lateral  pairs  arise  from  the  ninth  sternum,  the  dorsal 
pair  from  its  cephalic  margin  and  the  lateral  pair  from  its  caudal 
margin. 

The  ventral  valves  arising  from  the  eighth  sternum  are  two  long, 
slender  valves,  tapering  caudad  and  sharply  pointed  at  the  apices. 
They  are  ventral  in  position  and  inclose  the  dorsal  valves.  The 
ventral  and  dorsal  valves  of  each  side  are  joined  one  to  the  other 
by  a  tongue-and-groove  arrangement,  the  tongue  being  on  the  dor- 
sal valve  and  the  groove  on  the  ventral. 

The  dorsal  valves  are  innermost  of  the  three  pairs  and  are  dorsal 
in  location.  They  consist  of  a  narrow,  curved,  rodlike  base,  and  a 
broader,  flattened  apical  part  which  bears  teeth  along  its  dorsal 
edge.  The  apex  is  sharply  pointed  and  bears  very  fine  teeth  on 
both  dorsal  and  ventral  edges.  There  are  from  seventeen  to  nine- 
teen large  teeth  on  the  dorsal  edge. 

The  lateral  valves  are  the  outermost  of  the  three  and  enfold  the 
other  two  pairs.  They  are  broad,  flat,  and  somewhat  concave  on 
their  inner  surfaces.  They  are  not  capable  of  extension  as  are  the 
other  two  pairs,  for  in  addition  to  being  attached  basally  to  the 
ninth  sternum,  they  are  also  attached  to  the  pygofers  for  about  one- 
half  their  length.    Their  apical  half  is  free. 

Nymphal  Genital  Appendages. 

(Plate  XIX,  figs.   1,  3,  5,  7,  9,  11.) 

In  the  first  and  second  instars  two  pairs  of  very  small  chitinous 
pockets  may  be  seen,  one  projecting  caudad  from  the  eighth  sternum 
and  the  other  from  the  ninth.  In  the  first  pair  develop  the  ventral 
valves  of  the  adult  ovipositor,  and  in  the  second  the  dorsal  valves. 
In  these  two  instars  the  apices  of  the  ventral  pockets  are  at  a  dis- 
tance from  the  bases  of  the  dorsal  pockets.  The  dorsal  pockets, 
extending  caudad  for  about  one-third  the  length  of  the  ninth  seg- 
ment, are  also  short. 

In  the  third  instar  the  ventral  pockets  have  increased  in  length 
and  width.    They  are  still  short  and  broad.    Their  broadly  rounded 


Hackman:    Cicadella  Hieroglyphica.  199 

apices  overlap  the  bases  uf  the  dorsal  pockets.  The  narrower, 
longer  dorsal  pockets  extend  caudad  for  about  one-half  the  length 
of  the  ninth  segment.  In  this  instar  the  lateral  pockets  first  appear. 
They  lie  laterad  of  the  dorsal  pockets,  are  slightly  curved  and  some- 
what narrower  and  shorter  than  these. 

In  the  fourth  instar  the  ventral  pockets  are  longer  than  in  the 
preceding  instar,  and  are  now  more  than  two-thirds  the  length 
of  the  dorsal  pockets.  The  ventral,  still  the  longer  of  the  two  pairs, 
are  about  the  same  distance  from  the  tip  of  the  ninth  segment. 
They  are  beginning  to  be  enfolded  by  the  lateral  pockets,  which 
have  broadened  and  are  now  somewhat  concave  on  the  inner  sur- 
face. The  lateral  pockets  are  longer  than  the  ventral  pockets,  but 
shorter  than  the  dorsal. 

In  the  fifth  instar  the  three  pairs  of  pockets  are  rather  darkly 
chitinized.  The  ventral  pockets  are  broad  at  the  base  and  taper 
gradually  to  narrowly  rounded  apices.  They  are  the  shortest  of 
the  three  pairs,  though  nearly  as  long  as  the  dorsal  pair.  Only  the 
extreme  apices  of  the  dorsal  pockets  are  visible,  their  basal  portion 
being  hidden  by  the  broader  ventral  pockets.  They  are  still  longer 
than  the  ventral  pockets,  but  are  now  exceeded  in  length  by  the 
lateral  pockets.  These  are  somewhat  spoonlike  and  have  more  com- 
pletely enfolded  the  dorsal  pockets. 


200   -  The  University  Science  Bulletin. 


PLATE  XVI. 

1.  First  instar. 

2.  Second  instar. 

3.  Third  instar. 

4.  Fourth  instar. 

5.  Fifth  instar. 

6.  Adult. 


Hackman:    Cicadklla  Hip:roglyphica. 


201 


PLATE  XVI. 


■r^.vv".-.-,— ^ 

k. 

.^..,.,J 

p'.^"'-"'.'"'"^' 

1 

1 

t 

J 

T*^ 

1^..., 

J 

C^ 

"^"Z^ 

L, 

.,.,.■,  ^,.V.^-'.,'».t.'<>.'^ 

C , 

.....„..i 

r      5 

\    -/ 

^    * 

6 


202  The  University  Science  Bulletin. 


PLATE  XYH. 

1.  Egg  enlarged. 

2.  Leaf  with  two  egg  masses. 

3.  Leaf  showing  effect  of  ovii)ositioii. 

4.  Twig  showing  effect  of  oviposition. 


Hackman:    Ckadkli.a  Hierchjlyphica. 


203 


PLATE  XVII. 


204  The  University  Science  Bulletin. 


PLATE  XVIII. 

1.  Abdomen  of  adult  female. 

2.  First  sternite  of  female  abdomen. 

3.  Second  sternite  of  female  abdomen. 

4.  Ventral  valve  of  ovipositor  attached  to  eighth  sternum. 

5.  Doi-sal  valves  (upper)  and  lateral  valves  (lower),  showing  at'acl.int iit 
ninth  sternum. 

6.  Terminal  segments  of  female  abdomen,  showing  ovipositor. 

7.  First  sternite  of  male  abdomen. 

8.  Second  sternite  of  male  abdomen. 

9.  Abdomen  of  adult  male. 

10.  Cephalic  view  of  main  body  of  cedagus. 

11.  Caudal  view  of  main  body  of  oedagus. 

12.  CEdagus,  showing  attachment  of  ejaculatory  duct. 

13.  Styles  and  connective. 

14.  Ventral  valves,  dorsal  aspect,  showing  attachment  of  styles. 

15.  Male  genitalia. 


HacKMAN:     ('l(AI)KLLA    II 1  KKOdl.VI' 1 1 ICA. 


205 


PLATE  XVIII. 


%.=^ 


206  The  University  Science  Bulletin. 


PLATE  XIX. 

Terminal.  Segments  of  Abdomen; 

1.  Female,  first  instar. 

2.  Male,  first  instar. 

3.  Female,  second  instar. 

4.  Male,  second  instar. 

5.  Female,  third  instar. 

6.  Male,  third  instar. 

7.  Female,  fourth  instar. 

8.  Male,  fourth  instar. 

9.  Female,  fifth  instar. 

10.  Male,  fifth  instar. 

11.  Female  adult. 

12.  Male  adult. 


Hackman:    Cicadella  Hieroglyphica. 


207 


PLATE  XIX. 


208 


The  University  Science  Bulletin. 


PLATE  XX. 


Development  of  Male  Internal  Genitall\: 

1.  First  instar. 

2.  Second  instar. 

3.  Third  instar. 

4.  Fourth  instar. 

5.  Fifth  instar. 

6.  Fifth  instar,  phase  1,  dorsal  view. 

7.  Fifth  instar,  phase  1,  ventral  view. 

8.  Fifth  instar,  phase  2,  dorsal  view. 

9.  Fifth  instar,  phase  2,  ventral  view. 

10.  Fifth  instar,  phase  3,  dorsal  view. 

11.  Fifth  instar,  phase  3,  ventral  view. 

12.  Fifth  instar,  phase  3,  lateral  (right)  view  of  oedagus. 

13.  Fifth  instar,  phase  3,  lateral  (left)  view  of  cedagus. 

14.  Fifth  instar,  phase  4,  ventral  view  of  cedagus. 

15.  Fifth  instar,  phase  4,  lateral  (right)  view  of  oedagus. 

16.  Fifth  instar,  phase  4,  lateral  (left)  view  of  cedagus. 

17.  Fifth  instar,  phase  4,  lateral  (right)  view  of  oedagus. 

18.  Fifth  instar,  phase  4,  relative  position  of  plates,  styles  and  oedagus. 


Hackman:    Cicadella  Hieroolvphka, 


209 


PLATE  XX. 


3 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  8— October,  1922. 

(Whole  Series.  Vol.  XXIV,  No.    8.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 
Ovipositors  of  Cicadellide  (Homoptera) P.  A.  Readio. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE,  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


TABLE  OF  CONTENTS. 

PAGE 

Introduction 217 

The  abdomen  of  the  female 218 

The  ovipositor 220 

Taxonomic  use  of  the  ovipositor 222 

Descriptions  of  the  ovipositors  of  the  genera  and  species 224 

Conclusions 264 

Plates 267 

Bibliography 265 

Index 293 

(215) 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  8. 


The  Ovipositors  of  the  CicadelUdae  (Homoptera) . 

By  PHILIP  A.   READIO. 

.•^iihniitted  to  the  department  of  entomology  and  to  the  siaduHte  faculty  of  the  University 
of  Kansas  in  partial  fulfillment  of  the  requirements  for  the  degree  of  master  of  science, 
May   1.'),   1922. 

INTRODUCTION. 

THE  primary  purpose  of  this  paper  is  to  determine  whether  or 
not  the  ovipositors  of  the  Cicadelhclse  possess  characters  of 
value  in  classification,  and  if  so,  of  how  much  value  these  characters 
are  and  how  accessible  they  are  to  the  general  worker.  To  Prof. 
Paul  B.  Lawson  belongs  the  credit  for  suggesting  the  paper.  In  the 
same  group  he  has  recently  found  that  the  male  genitalia  are  of 
much  value  in  classification,  and  the  possibility  of  equal  value  in 
the  female  genitalia  occurred  to  him.  Miss  Itasca  Hilsman,  work- 
ing in  the  closely  related  family,  Cicadidse,  found  that  the  ovi- 
positors in  this  group  "afford  constant  and  ready  characters  which 
may  at  times  be  of  decided  value  to  him  (the  specialist)  in  the  de- 
termination of  closely  related  species."  Hence  it  was  natural  to  ex- 
pect that  equally  favorable  results  might  be  obtained  from  a  study 
of  the  ovipositors  of  the  Cicadellidse.  As  an  introduction  to  the 
taxonomic  part  of  the  paper,  a  morphological  study  of  the  abdomen 
of  the  female  and  the  structure  of  the  ovipositor  was  made. 

In  addition  to  the  reasons  already  stated,  there  are  certain  gen- 
eral considerations  which  would  lead  one  to  expect  such  an  investi- 
gation as  this  one  to  be  fruitful.  It  is  constant  characters  that  the 
taxonomist  is  in  search  of — characters  that  are  sufficiently  definite 
to  separate  closely  related  species,  and  yet  are  present  in  tlie  entire 
range  of  the  species.  Because  of  their  internal  position,  genitalia 
are  more  likely  to  be  constant  than  external  structures,  which  may 
vary  with  differences  of  environment.  The  constancy  of  the  use  to 
which  genitalia  are  put  also  makes  for  the  permanency  of  their 
structure  and  their  usefulness  as  taxonomic  characters. 


(217) 


218  The  University  Science  Bulletin. 

The  material  necessary  for  this  study  was  obtained  from  the 
duplicate  collection  of  the  University  of  Kansas  entomological  mu- 
seum. A  representative  number  of  ovipositors  from  each  of  the 
subfamilies,  excepting  the  Paropinse,  was  examined,  and  an  attempt 
made  to  examine  as  many  different  genera  and  as  many  species  in 
each  genus  as  possible.  In  all,  representatives  of  forty-eight  genera 
and  ninety-two  species  were  examined  and  figured. 

The  writer  wishes  to  express  to  Prof.  S.  J.  Hunter  his  appreciation 
of  the  sanction  given  to  this  work,  and  of  the  necessary  time  and 
materials  so  kindly  placed  at  his  disposal.  To  Prof.  P.  B.  Lawson 
many  thanks  are  due,  both  for  the  conception  of  the  nature  of  the 
problem  and  for  the  direction  of  the  work  to  its  completion.  It 
was  under  the  able  direction  of  Prof.  P.  W.  Claassen,  of  Cornell 
University,  that  the  photography  was  done.  To  Prof.  H.  B.  Hunger- 
ford,  Miss  Lucy  Hackman,  Miss  Kathleen  Doering  and  Mr.  Robert 
Guntert  the  writer  wishes  to  express  his  appreciation  for  their  kind 
help,  suggestions  and  criticisms. 

THE  ABDOMEN  OF  THE  FEMALE. 

(PI.  XXI,  figs.  1-3.) 

For  a  study  of  the  abdomen  of  the  female  leaf  hopper,  dried  speci- 
mens of  Oncoinetopia  lateralis  (Fabricius)  were  used.  These  were 
soaked  in  caustic  potash,  ten  per  cent,  for  twenty-four  hours,  and 
drawings  and  descriptions  made  from  the  cleared  specimens. 

The  abdomen  joins  the  thorax  broadly,  bulges  slightly  in  the  mid- 
dle, and,  from  a  point  a  little  past  the  middle,  tapers  both  in  width 
and  in  height  to  a  blunt,  caudal  point.  A  cross  section  has  a  general 
semicircular  outline.  The  tergum  is  arched,  appears  dorsally  and 
laterally,  and  forms  the  rounded  portion  of  the  semicircle,  while  the 
pleura  and  sternum  are  flat,  and  form  the  fiat,  ventral  portion.  The 
lateral  edges  of  the  sternum  are  bent  slightly  dorsad  at  their  union 
with  the  pleura.  The  pleura  and  tergum  unite  at  a  distinct  but 
slightly  rounded  angle. 

Eleven  segments  can  be  accounted  for  in  the  abdomen.  Of  these, 
segments  one  and  two  lack  distinct  pleura,  and  segments  eight  to 
eleven  are  variously  modified  as  described. 

The  tergum  of  the  first  segment  is  partly  membranous  and  partly 
chitinized.  Cephalad,  at  its  junction  with  the  metathorax,  it  is  en- 
tirely membranous.  Caudad,  at  its  junction  with  the  second  ab- 
dominal segment,  is  a  narrow,  linear,  chitinized  sclerite  which  bears 
laterally  two  inwardly  projecting  processes  along  its  cephalic  border. 


Readio:    Ovipositors  of  Cicadellid.f..  219 

Between  the  inwardly  projecting  processes  is  a  round,  small,  chi- 
tinized  piece.  The  pleura  are  entirely  membranous,  and  in  the 
pleural  membranes,  opening  ventrally,  are  the  spiracles  of  the  first 
segment.  These  are  larger  than  those  of  the  other  abdominal  seg- 
ments. The  sternum  consists  of  a  curved,  lightly  chitinized  caudal 
sclerite  extending  for  nearly  the  entire  width  of  the  segment,  and  of 
a  membranous  cephalic  portion  connecting  with  the  metatliorax. 
The  chitinized  portion  has  along  its  cephalic  border  two  pointed 
projections  which  point  mesad. 

The  tergite  of  the  second  segment  consists  of  a  simple,  linear, 
chitinized  sclerite  which  extends  nearly  the  entire  width  of  the  seg- 
ment. It  is  about  one-third  the  length  of  the  third  tergite.  The 
pleura  are  represented  only  by  pleural  membranes  in  which  the 
spiracles  of  this  segment  are  located.  The  spiracles  open  dorsally 
and  appear  to  be  in  the  membrane  laterad  of  the  second  tergite. 
The  sternum  of  segment  two  is  a  curved,  chitinous  piece  extending 
the  entire  width  of  the  segment. 

Segments  three,  four,  five  and  six  have  distinct  and  complete 
terga,  sterna  and  pleura.  The  terga  are  arched  and  form  the  dorsal 
and  lateral  surfaces.  The  ventral  surface  is  formed  by  the  pleura, 
which  are  subrectangular  and  longer  than  wide,  and  by  the  sterna, 
which  are  also  subrectangular,  but  wider  than  long.  Each  pleuron 
bears  a  spiracle  in  a  cephalomesal  position.  The  fourth  segment 
is  slightly  longer  than  the  third,  and  the  fifth  and  sixth  are  about  the 
same  length  and  longer  than  the  fourth. 

Segment  seven  is  also  complete.  The  tergum  and  pleura  are  the 
same  as  in  the  preceding  segments.  The  sternum  forms  the  sub- 
genital  plate  and  is  commonly  called  the  last  ventral  segment.  Its 
lateral  margins  converge  caudad  and  its  caudal  border  bears  a  broad 
but  shallow  indentation  along  its  mesal  half.  Dorsad  of  the  sternum 
of  segment  seven  is  an  invaginated  pocket  in  which  the  bases  of  the 
valves  of  the  ovipositor  are  located.  Its  ventral  side  is  formed  by 
the  apical  portion  of  the  seventh  sternum  and  a  membrane  arising 
from  the  dorsal  surface  of  this  sclerite  and  extending  cephalad  to 
its  base.  The  dorsal  side  of  the  pocket  is  formed  by  a  membrane 
which  is  continuous  with  the  membrane  of  the  ventral  side,  and 
which  extends  caudad  to  the  eighth  sterna  with  which  it  connects. 
The  greater  part  of  this  dorsal  membrane  is  strongly  chitinized,  but 
medially  it  is  apparently  divided  by  a  narrow,  clear  line  which  ex- 
pands apically  into  the  entirely  membranous  apex  connecting  with 


220  The  University  Science  Bulletin. 

the  eighth  sterna.  Laterally  this  pocket  is  bounded  by  the  continu- 
ation of  the  dorsal  membrane  to  the  seventh  and  eighth  pleura.  The 
anterior  portions  of  these  lateral  membranes  are  strongly  chitinized 
and  curve  mesad  ventrally. 

The  terminal  segments  of  the  female  abdomen  bear  the  external 
organs  of  reproduction  and  are  modified  for  this  purpose.  Segment 
eight  i^  represented  externally  only  by  the  tergum  and  the  pleura. 
The  tergum  is  essentially  the  same  as  in  the  preceding  segments, 
but  narrows  slightly  caudad.  The  pleura  narrow  caudad  and  are 
triangular  in  shape.  They  include  the  ventral  cephalic  portion  of 
the  ninth  segment  between  them  and  bear  the  last  pair  of  spiracles. 
Externally  segment  nine  consists  only  of  a  tergum,  commonly  called 
the  pygofer,  which  almost  completely  encircles  the  abdomen,  leav- 
ing only  a  ventral  groove  in  w'hich  the  ovipositor  is  folded.  Ven- 
trally it  extends  cephalad,  ending  in  two  obtuse  points  between  the 
eighth  pleurites.  Tlie  caudal  portion  encircles  nearly  the  whole  seg- 
ment and  tapers  towards  its  caudal  end. 

The  anal  tube  is  composed  of  segments  ten  and  eleven.  Segment 
ten  arises  from  the  narrowed  apex  of  segment  nine  and  is  short  and 
tubular.  It  connects  by  a  distinct  intersegmental  membrane  with 
segment  eleven,  which  is  also  tubular  and  can  be  telescoped  within 
segment  ten.  Segment  eleven  bears  the  telson,  the  extreme  apical 
portion  of  the  abdomen.  Along  the  dorsal  edge  of  the  telson  is  the 
anus,  an  opening  running  for  its  entire  length  and  guarded  by  hairs. 

There  are  eight  spiracles  in  the  abdomen  of  the  female  located  in 
the  first  eight  segments  as  described. 

Along  the  ventral  side  of  the  abdomen  there  occur  many  short, 
fine  hairs,  which  in  all  cases  point  backward.  The  hairs  on  the  ninth 
segment,  or  pygofer,  are  somewhat  longer  than  those  on  the  other 
parts  of  the  abdomen. 

THE  OVIPOSITOR. 

(PL   XXI,  figs.    4-9.) 

This  description  was  made  from  the  ovipositor  of  Oncometopia 
lateralis  (Fabricius).  However,  in  the  many  other  species  of  leaf 
hoppers  examined,  the  fundamental  structure  of  the  ovipositor  was 
lound  to  be  the  same  as  for  the  species  here  described. 

The  ovipositor  of  the  leaf  hopper  is  fitted  for  sawing  slits  in  plant 
tissues  and  for  placing  eggs  within  these  slits.  When  not  in  use  it 
lies  in  the  groove  in  the  ventral  surface  of  the  ninth  segment.  The 
ovipositor  consists  of  three  pairs  of  valves  or  gonapophyses.     The 


Readio:    OviPCSiTORs  of  Cicadellid.e.  221 

inner  two  pairs  alone  t'uiictioii  in  sawing  slits  and  placing  eggs,  and 
the  outer  pair  act  as  guards  within  which  the  two  inner  pairs  lie 
when  not  in  use.  In  this  discussion  the  valves  are  numbered  I,  II 
and  111,  according  to  their  attachment,  the  most  cephalic  in  attach- 
ment being  numbered  I. 

Valve  I  is  the  outer  and  more  ventral  of  the  two  inner  pairs  of 
\'alves.  Its  chief  attachment  is  to  the  eighth  sternum,  which  is 
divided  and  appears  as  two  subrectangular,  heavily  chitinized 
sclerites,  for  the  most  part  hidden  in  the  pocket  above  the  seventh 
sternum.  Near  its  base,  a  narrow,  curved  rod  leaves  its  dorsal  mar- 
gin and  attaches  basally  to  a  small,  triangular  sclerite  on  the  dorsal 
side  of  the  cephalic  end  of  the  pygofer.  This  small  triangular  scle- 
rite is  possibly  the  ninth  pleurite.  Upon  attempting  to  separate  the 
narrow,  rodlike  portion  from  the  rest  of  the  valve,  a  featherlike 
structure,  consisting  of  a  middle  portion  formed  by  the  continua- 
tion of  the  attaching  rod  and  membranous  side  plates,  can  be  drawn 
out.  However,  except  basally,  this  featherlike  structure  is  en- 
tirely incorporated  into  the  main  part  of  the  valve,  both  structurally 
and  functionally.  The  main  part  of  the  valve  is  relatively  broad 
basally,  narrows  slightly  beyond  the  base,  broadens  again  towards 
the  middle,  and  narrows  to  a  sharp,  apical  point.  A  heavily  chitin- 
ized rod  strengthens  the  ventral  portion  of  this  valve.  The  rod 
is  comparatively  large  basally,  narrows  in  the  middle  portion,  and 
is  absent  in  the  apical  third.  Along  the  entire  dorsal  border  and 
along  the  ventral  apical  border  are  diagonal  rows  of  scalelike  proc- 
esses. Along  its  mesal  surface,  just  dorsad  of  the  chitinized  rod,  is 
a  mushroom-shaped  groove  into  which  a  complementary  tongue  on 
the  outer  surface  of  valve  II  fits.  This  permits  independent  sliding 
of  the  two  valves,  but  not  complete  separation.  Basally  the  ven- 
tral edges  of  this  vah-e  turn  mesad,  and  on  the  dorsal  side  of  this  in- 
turned  portion  is  a  less  heavily  chitinized  projection.  This  projec- 
tion in  the  one  valve  bears  a  tongue,  and  in  the  other  a  groove,  unit- 
ing the  two  basally.  For  the  greater  part  of  their  length,  however, 
they  are  independent. 

Valve  II  is  the  inner  and  the  more  dorsal  of  the  two  inner  pairs 
of  valves.  It  is  attached  to  tlie  cephalic  end  of  a  small  sclerite, 
which  is  attached  to  the  pygofer.  This  sclerite  is  ventral  in  posi- 
tion and  possibly  represents  the  ninth  sternite.  This  valve  is  narrow 
and  rodlike  at  the  base,  but  for  its  greater  length  is  broad  and  flat. 
It  also  possesses  a  chitinized  strengthening  rod  whicli  extends  nearly 
to  its  apex.  On  the  outer  surface  of  the  valve,  just  ventrad  of  the 
rod,  is  a  tongue  which  fits  into  the  groove  in  valve  I.    The  dorsal 


222  The  University  Science  Bulletin. 

edge  usually  bears  teeth,  which  may  extend  for  its  entire  length,  be 
confined  to  the  apical  half,  or  appear  only  at  the  tip.  The  teeth 
are  of  various  shapes  in  the  different  species,  and  may  or  may  not 
in  turn  bear  secondary  teeth.  The  pointed  tip  is  usually  notched 
with  small  teeth,  and  these  may  appear  on  either  the  dorsal  or  ven- 
tral or  both  sides.  From  the  interior  of  the  valve  certain  ductlike 
structures  lead  to  the  dorsal  and  apical  teeth.  The  function  of  these 
ducts  is  unknown.  This  pair  of  valves  is  usually  united  one  to  the 
other  basally,  sometimes  by  a  heavy  chitinized  connection,  some- 
times only  by  a  membrane,  but  not  by  a  tongue  and  groove. 

Valve  III  is  the  most  caudal  in  attachment,  being  attached  to  the 
caudal  end  of  the  same  sclerite  to  which  valve  II  is  attached.  There 
is  also  a  membranous  connection  between  this  valve  and  the  ventral 
side  of  the  pygofer  for  about  h;ilf  the  length  of  the  valve.  The 
proximal  half  of  the  valve  is  narrow  and  the  apical  half  broad  and 
spoonlike,  being  somewhat  concave  on  its  inner  surface.  It  is 
bluntly  rounded  at  the  apex.  This  pair  of  valves,  between  which 
there  is  no  connection,  forms  a  protecting  sheath  in  which  the  ovi- 
positor is  completely  encased  when  not  in  use. 

Oviposition  of  Cicadella  hieroglyphica  (Say)  has  been  observed 
and  the  use  of  the  ovipositor  noted.  Valve  III  remains  in  its  posi- 
tion in  the  groove  of  segment  nine  and  takes  no  part  in  the  opera- 
tion. Valves  I  and  II,  closely  appressed  and  appearing  as  a  single 
structure,  are  extruded  from  their  resting  place  and  form  the  func- 
tional part  of  the  ovipositor.  The  ovipositor  is  first  held  vertically 
and  its  tip  inserted  under  the  epidermis  of  the  leaf.  It  is  then 
pushed  in  to  nearly  its  full  length,  and  now  is  in  a  horizontal  posi- 
tion, the  flat  surface  of  the  ovipositor  being  parallel  with  the  flat 
surface  of  the  leaf.  The  teeth  on  the  dorsal  edge  of  the  ovipositor 
are  now  turned  cephalad.  The  ovipositor  is  pushed  backward  and 
forward  with  a  sawlike  motion  until  the  chamber  is  large  enough  for 
the  reception  of  the  egg.  The  egg  passes  out  between  the  valves  of 
the  ovipositor  and  into  the  chamber,  and  when  in  place,  the  oviposi- 
tor is  withdrawn  and  folded  into  its  resting  place. 

* 

TAXONOMIC  USE  OF  THE  OVIPOSITOR. 

In  the  search  for  characters  of  taxonomic  value  in  the  ovipositor, 
valve  II  was  at  once  hit  upon  as  being  the  structure  most  likely  to 
possess  useful  characters.  It  varies  in  different  species  in  regard  to 
size,  shape,  number  and  shape  of  primary  teeth,  number  and  shape 
of  the  secondary  teeth  borne  by  the  primary  teeth,  characters  of  the 
tip,  and  in  the  number  and  arrangement  of  the  ducts.     That  these 


Readio:    Ovipositors  of  Cicadp:llid^.  223 

characters  are  constant  within  (he  species  has  been  proven  by  ex- 
amining a  wide  range  of  in(h\-i(hials  within  the  species  and  finding 
that  the  \-ariation  was  negligible.  C'onst'ciuently,  for  the  taxononiic 
part  of  this  paper,  valve  II  is  used  exclusively. 

The  technique  used  in  mounting  valve  II  for  study  is  simple.  The 
tip  of  the  abdomen  bearing  the  oN'ipositor  is  broken  off  from  the 
dried  specimen,  soaked  in  ten  i:>er  cent  caustic  potash  for  twenty- 
four  hours,  washed  in  water  for  a  few  minutes,  valve  II  and  other  de- 
sired parts  dissected  out  under  a  binocular  microscope,  dehydrated 
in  ninety-five  per  cent  alcohol  for  five  minutes,  cleared  in  xylol  for 
five  minutes,  and  mounted  in  balsam  on  a  microscope  slide.  It  is 
well  to  give  the  slide  and  the  specimen  from  which  the  slide  w^as 
made  a  corresponding  number,  so  that  any  necessary  checks  may 
be  made.  This  method  gives  a  permanent  mount,  which  may  be 
studied  at  the  convenience  of  the  worker. 

Several  methods  of  figuring  the  ovipositor  were  used.  The  first 
was  that  of  drawing  the  desired  valve  free-hand  with  the  aid  of  a 
micrometer  eyepiece  divided  into  squares,  which  correspond  to 
squares  on  the  drawing  paper.  This  is  a  satisfactory  method,  but 
somew^hat  more  laborious  than  the  methods  later  used.  The  second 
was  to  draw  with  the  aid  of  the  camera  lucida.  The  particular 
equipment  available  did  not  give  satisfactory  results  at  all  times, 
but  there  is  no  doubt  but  what  this  method  could  be  used  with 
satisfaction.  The  greatest  degree  of  success  was  obtained  by  the 
use  of  the  Edinger  drawing  apparatus.  This  apparatus  projects  the 
desired  image  upon  the  drawing  paper,  and  the  figuring  consists  only 
of  tracing  in  the  image.  Photography  w'as  tried  and  found  to  be  a 
very  successful  means  of  reproducing  the  desired  image.  A  camera 
fitted  for  ordinary  scientific  work  was  used.  It  w^as  turned  to  a  ver- 
tical position  and  a  lens  board  fitted  with  a  black,  light-proof  sleeve 
was  inserted  in  the  front  lens-board  holder.  The  sleeve  was  lowered 
over  the  ocular  of  a  compound  microscope,  and  focusing  for  desired 
size  and  definition  of  image  accomplished  by  a  combination  of 
microscope  and  camera  adjustments.  Various  substitutions  in  ob- 
jectives and  oculars  were  necessary  to  meet  all  conditions.  Illumi- 
nation was  furnished  by  an  ordinary  substage  light,  and  exposures 
were  made  by  the  switching  off  and  on  of  this  lamp.  Undoubtedly 
much  better  results  could  be  obtained  with  the  aid  of  special  photo- 
micrographic  equipment. 

Descriptions  of  the  ovipositors  of  the  generji  and  species  of  the 
various  subfamilies  follow: 


224  The  University  Science  Bulletin. 

Subfamily  BYTHOSOPIN.^  (Dohrn). 
Agalliopsis  novella  (Say). 

(PI.  XXII,  fig.   2.:    pi.   XXV,  fig.   1.) 

Length,  1.1  mm.;  greatest  width,  0.11  mm.  Apical  toothed  half  only 
slightly  wider  than  basal  half;  distinctly  curved,  tip  rather  gradually  narrowed, 
extreme  apex  somewhat  pointed,  chitinization  light;  strengthening  rod  ex- 
tends caudad  as  far  as  twenty-first  or  twenty-second  dorsal  tooth  from  apex. 
Toothed  area  on  doi-sal  edge  occupies  slightly  less  than  the  apical  half,  teeth 
sixty-three  to  sixty-five  in  number,  very  small,  angular,  pointed,  rather  regu- 
lar in  size,  shape  and  spacing;  a  few  double  teeth  present,  no  secondary  teeth; 
the  tip  is  notched  on  both  edges  with  small  teeth,  practically  continuous 
around  the  tip,  twenty  to  twenty-two  on  \'entral  edge;  area  of  ducts  in- 
conspicuous, a  few  ducts  faintly  visible  for  their  entire  length,  but  for  the 
most  part  are  invisible  except  for  their  apices  and  circular  openings,  open 
along  the  ventral  apical  edge  (seven),  along  the  dorsal  edge  of  the  toothed 
area,  and  in  the  basal  region;  the  two  valves  of  the  pair  are  joined  one  to 
the  other  by  a  narrow,  elongate,  chitinized  connection  present  on  the  dorsal 
edge  of  the  basal  region  at  a  point  about  two-fifths  the  length. 

Aceratagallia  uhleri  (Van  Duzee). 

(PI.  XXV,  fig.  2.) 

Length.  1  mm.;  greatest  width,  .07  mm.  Apical  half  very  little  if  any 
wider  than  basal  half;  .distinctly  curved,  tip  gradually  narrowed,  extreme 
apex  rather  pointed,  chitinization  moderately  light;  strengthening  rod  ex- 
tends caudad  as  far  as  twenty-second  dorsal  tooth  from  apex.  Toothed  area 
on  dorsal  edge  occupies  the  apical  half  of  the  length;  teeth  about  ninety 
in  number,  very  small,  angular,  pjointed,  fairly  regular  as  to  size,  shape  and 
spacing,  larger  apically;  bear  no  secondary  teeth;  the  tip  is  notched  with  small 
teeth  on  both  edges,  continuous  around  the  tip,  about  twenty-two  on  the 
^•entral  edge;  area  of  ducts  conspicuous,  ducts  easily  visible  for  entire  length; 
open  along  ventral  apical'  edge  (five  to  six),  along  dorsal  edge  of  toothed 
area,  and  in  basal  region;  the  two  valves  of  the  pair  are  joined  one  to  the 
other  by  an  elongate,  naiTOW,  chitinized  connection  present  on  the  dorsal  edge 
at  a  point  about  one-third  the  length. 

This  ovipositor  is  similar  to  that  of  Agalliopsis  novella  (Say)  in 
general  appearance. 

Genus  Idiocerus  Lewis. 

The  ovipositors  of  seven  species  of  this  genus  have  been  examined 
and  found  to  be  generally  similar.  In  each  case  the  ovipositor  is 
about  the  same  width  for  the  entire  length  and  is  only  slightly 
curved.  Examples  of  light,  medium  and  heavy  chitinization  are 
found.  The  valve  may  be  toothed  along  its  dorsal  edge  from  a 
fourth  to  a  little  more  than  a  half  its  length  apically.  The  teeth 
in  the  specimens  examined  are  ten  to  thirty  in  number,  medium  to 
large  in  size,  rounded,  evenly  spaced,  and  may  or  may  not  bear 
secondary  teeth.     The  ducts  may  appear  ductlike,  granular,  or  be 


Readio:    Ovipositors  of  Cicadellid.e.  225 

invisible.  The  apex  may  or  may  not  bear  small  teeth  along  one 
or  both  edges.  A  ohitinous  connection  is  present  on  the  dorsal  edge 
of  the  basal  area  in  some  species. 

Idioccrus  ncrvatus  Van  Duzee. 

(PL  XXV,  fig.  9.) 

Length,  1.4  nun.;  greatest  width,  0.12  nun.  Apical  portion  slightly  wider 
than  basal  portion;  shghtly  curved,  tip  greatly  narrowed  with  the  extreme 
apex  bluntly  rounded,  veiy  lightly  chitinized;  strengthening  rod  extends 
caudad  as  far  a^;  eleventh  doi-sal  tooth.  Toothed  area  on  dorsal  edge  occupies 
the  apical  two-fifths  of  the  length;  teeth  fourteen  to  sixteen  in  number,  large, 
long,  rounded,  regular,  evenly  spaced,  though  somewhat  farther  apart  basally; 
bear  no  secondary  teeth,  though  the  margin  is  irregularly  roughened;  the  tij) 
is  notched  on'  both  edges  with  small,  forward-pointing  teeth,  eight  appear 
ventrally,  one  to  three  dorsally;  area  of  ducts  inconspicuous,  circular  openings 
alone  visible;  of  these  four  open  ventrally,  three  apically,  and  many  dorsally. 
There  is  no  evidence  of  a  chitinous  connection. 

Idiocerus  pallidus  Fitch.  . 

(PI.  XXII,  fig.   1;    p!.   XXV,  fig.   5.) 

Length,  2.4  mm.;  greatest  witlth,  0.25  mm.  About  the  same  width  for 
entire  length;  slightly  curved,  tip  bluntly  rounded,  heavily  chitinized;  strength- 
ening rod  extends  caudad  as  far  as  ninth  dorsal  tooth.  Toothed  area  on 
doi-sal  edge  occupies  a  little  more  than  the  apical  third;  teeth  fourteen  in 
number,  large,  rounded,  regular,  evenly  spaced  except  basally,  where  they  are 
farther  apart;  secondary  teeth  only  on  last  eight  primary  teeth,  these  very 
small  and  indistinct,  especially  on  those  farthest  from  the  tip.  where  they  can 
be  seen  only  with  the  aid  of  high-power  magnification;  the  tip  is  notched  with 
small,  regular  teeth  on  both  dorsal  and  ventral  edges;  area  of  ducts  con- 
spicuous, ducts  elongate,  large,  straight;  six  open  along  ventral  apical  edge, 
one  in  extreme  apex,  and  many  along  the  dorsal  edge.  A  rather  indistinct 
chitinous  connection  is  present  on  the  doi-sal  edge  of  the  basal  area  and  joins 
the  two  valves  of  the  pair. 

Idiocerus  diizeei  Provancher. 

(PI.  XXV,  fig.  6.) 

Length.  2.6  mm.;  greatest  width,  0.34  mm.  Apical  portion  slightly  wider 
than  basal  portion;  slightly  curved,  tip  ends  in  somewhat  rounded,  obtuse 
point,  heavily  chitinized;  strengthening  rod  extends  caudad  as  far  as  sixth 
dorsal  tooth.  Toothed  area  on  dorsal  edge  occupies  a,  httle  less  than  the 
apical  half,  teeth  nine  to  ten  in  number,  large,  somewhat  angular,  regular, 
evenly  spaced,  but  larger  and  farther  apart  basally,  teeth  one  and  two  being 
veiy  large,  rounded,  and  heavily  chitinized;  all  the  primary  teeth  except 
teeth  one.  two  and  three  are  notched  with  small  secondary  teeth;  tip  is 
notched  with  many  small,  regular  teeth  on  both  dorsal  and  ventral  edges; 
ducts  conspicuous,  rather  small,  straight,  open  by  circular  openings;  six  open 
along  the  ventral  apical  edge,  one  in  extreme  apex,  and  many  along  the 
doi-sal  edge;  there  is  no  well-defined  chitinized  connection  between  the  two 
valves  of  the  pair. 


226  The  University  Science  Bulletin. 

Idiocerus  verticis  (Say). 

(PI.  XXV,  fig.  V.) 

Length,  1.55  mm.;  greatest  width,  0.15  mm.  Apical  portion  only  sHghtly 
wider  than  basal;  slightly  curv^ed,  tip  greatly  narrowed  with  extreme  apex 
rounded,  chitinization  light;  strengthening  rod  extends  caudad  as  far  as 
fourteenth  dorsal  tooth.  Toothed  area  on  dorsal  edge  occupies  a  little  more 
than  the  apical  third ;  teeth  twenty  in  number,  large,  rounded,  regular,  evenly 
spaced  except  basally;  bear  no  distinct  secondary  teeth,  though  the  margin 
is  irregularly  roughened  suggesting  secondary'  teeth;  the  tip  bears  ten  small 
caudad-pointing  teeth  along  its  ventral  edge;  the  ducts  are  inconspicuous, 
their  circular  openings  alone  being  visible;  of  these  apparently  two  open 
ventrally,  three  apicalty,  and  many  dorsally;  there  is  no  distinct  chitinized 
connection  between  the  two  valves  of  the  pair. 

Idiocerus  scurra  (Germar) . 

(PI.  XXV,  fig.  8.) 

Length,  222  mm.;  greatest  width,  0.28  mm.  About  the  same  width  for 
entire  length;  slightly  cui-ved,  tip  nan-owed  but  broadly  rounded  at  the  ex- 
treme apex,  rather  heavily  chitinized;  strengthening  rod  extends  caudad  as 
far  as  the  twenty-fifth,  dorsal  tooth.  Toothed  area  on  dorsal  edge  occupies 
a  little  more  than  the  apical  half;  teeth  thirty  in  number,  moderately  large, 
rounded,  regular,  evenly  spaced,  about  the  same  distance  apart  for  entire, 
length;  bear  no  secondary  teeth;  the  ventral  edge  of  the  tip  bears  ten  small 
teeth;  area  of  ducts  conspicuous,  granular  in  appearance,  circular  openings 
visible;  two  open  ventrally,  two  apically,  and  many  dorsally;  the  two  valves 
of  the  pair  are  joined  one  to  the  other  by  a  distinct,  elongate,  chitinized  con- 
nection present  on  the  dorsal  edge  of  the  basal  area. 

Idiocerus  ramentosus  (Uhler) . 

(PI.  XXV,  fig.  4.) 

Length,  1.96  mm.;  greatest  width,  0.17  mm.  About  the  same  width  for  en- 
tire length;  only  slightly  curved,  tip  greatly  narrowed  with  extreme  apex 
rounded,  moderately  heavy  chitinization;  strengthening  rod  extends  caudad 
as  far  as  eleventh  dorsal  tooth.  Toothed  area  on  dorsal  edge  occupies  a  little 
more  than  the  apical  third ;  teeth  eighteen  in  number,  large,  rounded,  generally 
regular  with  a  few  double  teeth,  evenly  spaced  but  farther  apart  basally;  bear 
no  secondary  teeth;  the  tip  bears  three  to  four  small,  inconspicuous  teeth 
along  its  ventral  edge;  area  of  ducts  conspicuous,  granular  in  appearance, 
circular  duct  openings  visible;  two  to  three  open  ventrally,  three  apically, 
and  many  doreally;,  the  two  halves  of  the  pair  joined  one  to  the  other  by 
an  elongate,  narrow,  chitinized  connection  present  on  the  dorsal  edge  of  the 
basal  area. 

Idiocerus  snowi  Gillette  and  Baker. 

(PI.  XXV,  fig.  3.) 

Length,  2.33  mm.;  greatest  width,  0.22  mm.  About  the  same  width  for 
entire  length;  slightly  curved,  tip  very  bluntly  rounded,  rather  heavily  chitin- 
ized; strengthening  rod  extends  caudad  as  far  as  eighth  dorsal  tooth. 
Toothed  area  on  dorsal  edge  occupies  the  apical  fourth;  teeth  ten  in  number, 


Readio:    Ovipositors  of  Cicadellid.e.  227 

medium  in.  size,  broad  ami  rounded,  regularly  spaced  except  basally,  where 
they  are  farther  apart;  bear  no  secondary  teeth;  no  teeth  present  at  the 
apex,  area  of  ducts  conspicuous,  granular  in  appearance,  circular  duct  open- 
ings visible,  none  open  ventrallj-,  six  apically,  and  many  dorsally;  the  two 
valves  of  the  pair  are  joined  one  to  the  other  by  an  elongate,  narrow,  chitin- 
ized  connection  present  on  the  doi-sal  edge  of  the  basal. area. 

Genus  Macropsis  Lewis. 
The  ovipositors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  very  similar.  They  are  long,  narrow  and  rodlike, 
about  the  same  width  for  entire  length,  slightly  curved,  and  narrow 
only  at  the  apex  to  an  obtuse  point.  The  two  valves  of  the  pair  are 
not  identical  as  to  length,  shape  of  tip,  and  teeth  found  at  tip.  The 
toothed  area  occupies  only  a  small  part  of  the  length  at  the  apex. 
The  teeth  are  few  in  number,  rather  large,  rounded,  and  may  or 
may  not  bear  secondary  teeth.  The  tip  is  notched  with  small  teeth 
in  both  species.  The  ducts  are  elongate,  conspicuous,  few  in  num- 
ber, and  located  only  apically. 

Macropsis  viridis  (Fitch). 

(PI.  XXVI,  fig.  1.) 

Length,  2.3  mm.;  greatest  width,  0.05  mm.  Very  long,  narrow,  rodlike, 
about  the  same  width  for  entire  length,  the  two  valves  of  the  pair  are  not 
identical  in  length,  shape  of  tip,  and  teeth  found  at  tip;  slighth'  curved,  nar- 
rows at  apex  to  obtuse  point,  rather  heavily  chitinized,  strengthening  rod 
extends  caudad  as  far  as  second  dorsal  tooth.  Toothed  area  occupies  but  a 
small  portion  at  the  apex,  the  longer  valve  bears  two  large  teeth  on  its  dorsal 
edge  and  eleven  small,  inconspicuous  teeth  along  its  ventral  edge,  the  shorter 
valve  bears  three  large  teeth  along  its  dorsal  edge  and  eleven  small  teeth 
along  its  ventral  edge,  none  of  these  teeth  bear  secondaiy  teeth;  the  extreme 
apex  of  the  longer  valve  is  entirely  devoid  of  teeth,  while  the  ventral  teeth 
of  the  shorter  valve  extend  to  the  apex,  ducts  are  conspicuous,  few  in  num- 
ber, elongate,  all  opening  apically,  two  to  four  openings  visible. 

Macropsis  suturalis  (Osborn  and  Ball). 

(PI.  XXVI,  fig.  2.) 

Length,  2.1  mm.;  greatest  width,  0.05  mm.  Very  long,  narrow,  rodlike, 
about  the  same  width  for  entire  length;  the  two  valves  of  the  pair  are  not 
identical,  but  differ  in  length,  shape  of  tip,  and  teeth  at  tip;  slightly  curved, 
narrows  at  tip  to  an  obtuse  point,  chitinization  moderately  heavy,  some- 
what lighter  than  in  M.  viridis;  strengthening  rod  extends  caudad  as  far  as 
fifth  dorsal  tooth.  Toothed  area  includes  only  the  apical  portion ;  the  longer 
valve  bears  on  its  dorsal  edge  two  large,  heavily  chitinized  teeth  at  the  tip 
and  three  large,  somewhat  less  heavily  chitinized  teeth  back  from  the  tip; 
on  its  ventral  edge  are  seventeen  small  teeth  which  are  continuous  around 
the  tip;  the  shorter  valve  bears  three  large,  heavily  chitinized  teeth  at  the 
tip,  three  less  heavily  chitinized  teeth  back  from  the  tip,  both  groups  being 


228  The  University  Science  Bulletin. 

on  the  dorsal  edge,  and  tliirteen  small  teeth  on  the  ventral  and  apical  edges; 
the  three  large  teeth  at  the  apex  of  the  shorter  valve  bear  small  secondary 
teeth;  the  ducts  are  conspicuous,  few  in  number,  all  located  at  the  apex,  six 
to  seven  openings  visible. 

Oncopsis  distinctus  (Van  Duzee). 

(PI.  XXVI,  fig.  3.) 

Length,  1.5  mm.;  greatest  width,  0.08  mm.  Long,  narrow,  rodlike,  about 
the  same  width  for  entire  length;  the  two  valves  of  the  pair  are  not  identical, 
but  differ  in  length,  shape  of  tip,  and  teeth  found  at  tip;  slightly  curved, 
narrow  at  apex  to  obtuse  point,  heavily  chitinized;  strengthening  rod  ex- 
tends caudad  as  far  as  the  second  dorsal  tooth.  Toothed  area  on  dorsal  edge 
occupies  only  the  apical  portion  of  the  valve;  the  longer  valve  bears  two  large, 
heavily  chitinized  teeth  on  the  dorsal  edge  and  fourteen  small,  inconspicuous 
teeth  on  the  ventral  edge,  and  the  shorter  valve  bears  two  large,  heavily 
chitinized  teeth  on  the  dorsal  edge  and  fourteen  small,  inconspicuous  teeth 
on  its  ventral  and  apical  edges;  none  of  the  teeth  bear  secondary  teeth;  the 
ducts  are  conspicuous,  elongate,  few  in  number;  five  to  six  open  apically, 
and  others  are  scattered  along  the  basal  rod  of  the  valve  and  open  dorsally. 

Genus  Bythoscopus  Germar. 
The,  ovipositors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  simiUar.  They  are  about  the  same  width  for  the 
entire  length;  slightly  though  distinctly  curved,  narrow  abrutly  near 
the  apex,  and  end  in  a  rounded  extreme  apex.  The  teeth  on  the 
dorsal  edge  occupy  about  a  third  of  the  apical  length,  are  numerous, 
small,  irregular,  and  bear  no  secondary  teeth.  There  are  teeth  on 
the  ventral  edge  of  tip  in  both  species.  The  ducts  are  conspicuous, 
elongate,  rather  few  in  number.  In  both  species  there  is  present  on 
the  dorsal  edge  about  midway  of  the  valve  an  elongate,  distinct, 
heavily  chitinized  connection  between  the  two  valves  of  the  pair. 

Bythoscopus  apicalis  (Osborn  and  Ball). 

(PI.    XXV,    fig.    10.) 

Length,  1.53  mm.;  greatest  width,  0.14  mm.  The  apical  portion  is  slightly 
wider  than  the  basal  portion,  distinctly  curved;  the  valve  narrows  from  a 
point  about  six-sevenths  of  its  entire  length  to  the  tip,  extreme  apex  rounded, 
chitinization  moderately  heavy;  strengthening  rod  extends  caudad  to  within 
six  dorsal  teeth  from  the  apex.  Toothed  area  on  the  dorsal  edge  occupies  a 
little  more  than  the  apical  third  of  the  length;  teeth  numerous,  very  small 
and  inconspicuous,  flatly  rounded,  irregular  in  shape  and  size,  unevenly 
spaced;  bear  no  secondary  teeth;  the  ventral  edge  of  the  tip  bears  a  few  in- 
conspicuous teeth;  ducts  conspicuous,  comparatively  few  in  number,  one  group 
opens  apically,  and  another  group  along  the  basal  rod  of  the  valve  about  mid- 
way; the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a  distinct, 
elongate,  chitinized  connection  present  on  the  dorsal  edge  about  midway  of 
the  valve,  this  connection  bearing  an  angular  tooth  near  its  caudal  extremity. 


Readio:    Ovipositors  of  Cicadellid.e.  229 

Bythoscoptis  miscellus  (Stal). 

(PI.  XXV,  fig.   11.) 

Length,  1.27  iniu.;  greatest  width,  0.08  mm.  About  the  same  width  for 
entire  length;  distinctlj^  curved,  tip  narrowed  abruptly  from  a  slight  promi- 
nence on  the  dorsal  edge  about  eight-ninths  of  the  length  to  a  broadly  rounded 
e.xtreme  apex;  chitinization  medium  heavj',  somewhat  lighter  than  in  B. 
apicnlis;  strengthening  rod  extends  caudad  as  far  as  fourteenth  doi-sal  tooth 
from  the  apex.  Toothed  area  on  dorsal  edge  occupies  a  little  less  than  the 
apical  third;  teeth  numerous,  small,  rather  sharjily  pointed,  irregular,  un- 
evenly si)aoed:  bear  no  secondary'  teeth;  the  tip  is  notched  with  teeth  which 
are  continuous  around  the  tip,  ten  to  eleven  on  the  ventral  edge;  ducts  con- 
spicuous, elongate,  few  in  number;  one  group  opens  apically,  another  at  the 
basal  end  of  the  toothed  area,  and  another  in  the  basal  rod  about  midway; 
the  tw^o  valves  of  the  pair  are  joined  one  to  the  other  by  a  distinct,  elongate, 
chitinized  connection  jiresent  on  the  dorsal  edge  about  midway. 

Subfamily  CICADELLINiE  Van  Duzee. 

Genus  Oncometopia  Stal.  ^ 

The  ovipositors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  simihir,  though  having  some  very  distinct  differ- 
ences. The  ovipositor  of  0.  iindata  more  closely  resembles  the 
ovipositor  of  Homalodisca  triquetra  than  it  does  the  ovipositor  of 
0.  lateralis.  In  the  ovipositors  of  this  genus  we  find  a  narrow, 
curved,  rodlike  basal  attachment,  and  a  broad,  fiat,  apical  shaft 
bearing  teeth  along  its  dorsal  edge,  and  a  preapical  prominence  on 
its  ventral  edge.  The  primary  teeth  in  each  case  bear  secondary 
teeth.  The  tip  is  notched  with  small  teeth  on  both  edges;  the  ducts 
are  conspicuous,  elongate,  numerous,  and  distinctly  curved. 

Oncometopia' undat a  (Fabricius) . 

(PI.  XXII,  fig.   3;    pi.  XXVI,  fig.   4.) 

Length  of  toothed  area,  L8  mm.;  greatest  width,  0.27  mm.  Narrow  and 
rodlike  at  base,  widens  beyond  base  into  flat,  broad  apical  shaft,  which  is 
about  the  same  width  for  entire  length  and  tapers  only  at  the  apex;  pre- 
apical prominence  present,  conspicuous,  broadly  rounded;  apical  shaft  slightly 
curved,  tip  broadly  rounded,  chitinization  medium;  strengthening  rod  extends 
caudad  as  far  as  last  doi-sal  tooth.  Toothed  area  on  dorsal  edge  occupies 
entire  length  of  broadened  shaft;  teeth  thirty-one  to  thirty-two  in  number, 
of  medium  size,  of  a  general  triangular  shape  with  the  caudal  side  much 
longer  than  the  cephalic  side;  apices  rounded,  regular,  evenly  spaced,  more 
heavily  chitinized  than  the  rest  of  the  valve;  possess  secondary  teeth  on  both 
caudal  and  cephalic  edges,  one  to  four  on  cephahc  edge  and  two  to  five  on 
caudal  edge,  those  on  the  caudal  edge  much  larger  than  those  on  the  cephalic 
edge;  tip  notched  with  small  teeth  on  both  dorsal  and  ventral  edges,  not  con- 
tinuous around  the  tip,  fifteen  teeth  present  on  ventral  edge  between  preapical 


230  The  University  Science  Bulletin. 

prominence  and  extreme  apex;  ducts  conspicuous,  decidedly  cui-ved,  numerous, 
most  numerous  apically;  open  apically  and  along  dorsal  edge  in  and  near 
teeth,  one  to  three  to  each  tooth. 

Oncometopia  lateralis  (Fabricius). 

(PI.  XXII,  fig.   4;   pi.  XXVI,  fig.   5.) 

Length  1.66  mm.;  greatest  width,  0.2  mm.  Narrow  and  rodlike  at  the  base, 
widens  beyond  base  into  broad,  flat  apical  shaft,  which  tapers  very  slightly 
toward  apex;  preapical  prominence  present,  rather  indistinct,  broadly  rounded; 
only  very  slightly  cui-ved  beyond  the  base,  tip  broadly  rounded,  chitinization 
medium,  strengthening  rod  extends  caudad  as  far  as  last  dorsal  tooth.  Toothed 
area  on  dorsal  edge  occupies  the  entire  length  of  the  apical  shaft;  teeth  eigh- 
teen to  twenty  in  number,  large,  subquadrate  in  shape  with  the  cephalic  side 
longer  than  the  caudal  and  bearing  a  smoothly  rounded  prominence,  regular, 
evenly  spaced,  more  heavily  chitinized  than  the  rest  of  the  valve  and  have 
the  appearance  of  being  set  into  the  rest  of  the  valve;  bear  secondary  teeth, 
seven  to  fifteen  on  the  broad  outer  edge  and  four  to  eight  on  the  cephalic 
side,  those  on  the  outer  edge  larger  than  those  on  the  cephalic  side;  the  tip 
is  notched  with  small  teeth  on  both  dorsal  and  ventral  edges,  not  continuous 
around  the  tip,  twenty-one  teeth  between  preapical  prominence  and  extreme 
apex;  ducts  conspicuous,  elongate,  distinctly  curved,  numerous,  most  numerous 
apically ;  open  apically,  along  the  dorsal  edge  in  and  near  teeth,  two  to  four  to 
each  tooth;  first  three  teeth  not  served  with  ducts. 

Homalodisca  triquetra  (Fabricus). 

(PI.  XXII,  fig.  5;   pi.  XXVI,  fig.  6.) 

Length,  2.6  mm.;  greatest  width,  0.33  mm.  Narrow  and  rodlike  at  base, 
widens  beyond  base  into  broad,  flat  portion,  which  is  about  the  same  width 
for  entire  length,  narrowing  only  at  apex;  preapical  prominence  present, 
distinct,  broadly  rounded;  uncurved  beyond  base,  extreme  apex  broadly 
rounded,  chitinization  medium;  strengthening  rod  extends  caudad  as  far  as 
third  dorsal  tooth  from  the  apex.  Toothed  area  on  dorsal  edge  occupies  the 
entire  length  of  the  apical  shaft;  teeth  forty-four  to  forty-five  in  number, 
medium  in  size,  of  a  general  triangular  shape  with  the  caudal  side  longer  than 
the  cephalic,  regular  in  size  and  shape,  evenly  spaced,  more  heavily  chitinized 
than  the  rest  of  the  valve ;  bear  secondary  teeth,  three  to  seven  on  the  cephalic 
edge  and  five  to  nine  on  the  caudal  edge;  those  on  the  caudal  edge  are  larger 
than  those  on  the  cephaUc  edge;  tip  notched  with  small  teeth  on  both  dorsal 
and  ventral  edges,  not  continuous  around  the  tip,  ducts  conspicuous,  numerous, 
slightly  curved,  most  numerous  apically;  open  apically  along  the  dorsal  edge 
in  and  near  teeth,  two  to  five  to  each  tooth;  the  first  six  teeth  not  served  by 
ducts. 

Aulacizes  irrorata  (Fabricius), 

(PI.  XXII,  fig.  6;   pi.  XXVI,  fig.  7.) 

Length,  2.9  mm.;  greatest  width,  0.44  mm.  Narrow  and  rodlike  at  the  base, 
widens  beyond  base  into  broad,  flat,  apical  shaft;  reaches  point  of  greatest 
width  about  two-fifths  of  length  and  from  that  point  tapers  gradually  to  the 
tip;   preapical  prominence  present,   conspicuous,   obtuse-angled;    not   curved 


Readio:    Ovipositors  of  Cicadellid.e.  231 

be5'ond  curved  basal  attachmont;  tip  bluntly  rounded,  chitinization  medium; 
strengthening  rod  extends  caudad  as  far  as  third  dorsal  tooth  from  apex. 
Toothed  area  on  dorsal  edge  occupies  the  entire  length  of  the  apical  shaft; 
teeth  forty-one  to  fortj'-thrce  in  number,  rather  small,  in  the  shape  of  a  flat- 
tened isosceles  triangle,  regular  in  size  and  shape,  evenly  spaced;  the  first 
twelve  teeth  bear  no  secondary  teeth;  the  teeth  caudad  of  the  first  twelve  teeth 
bear  four  to  ten  secondary  teeth  on  the  caudal  side;  tip  notched  with  small 
teeth  on  both  dorsal  and  ventral  edges,  not  continuous  aroimd  the  tip;  thirty- 
five  to  thirtj'-seven  teeth  on  ventral  edge  between  preapical  prominence  and 
extreme  apex;  ducts  conspicuous,  elongate,  rather  narrow,  curving,  numerous, 
most  numerous  apically;  open  apically,  along  dorsal  edge,  near  but  not  in 
teeth,  and  in  face  of  valve  back  from  edge;  the  first  twelve  teeth  are  not 
served  by  ducts. 

Genus  Cicadella  Latreille. 

The  ovipcsitors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  similar.  In  each  case  the  ovipositor  consists  of  a 
narrow,  curved  basal  attachment  and  a  wider  apical  shaft  which 
bears  teeth  along  its  dorsal  edge,  in  turn  bearing  secondary  teeth. 
A  preapical  prominence  is  present  in  one  species,  absent  in  the  other. 
The  tip  is  notched  on  both  dorsal  and  ventral  edges.  The  ducts 
are  rather  inconspicuous,  elongate,  rather  few  in  number,  and  dis- 
tinctly curved. 

Cicadella  hicroglyphica  (Say). 

(PI.  XXII,  fig.  7;    pi.  XXVI,  fig.  9.) 

Length,  1.66  mm.;  greatest  width,  0.13  mm.  Curved  basal  attachment  nar- 
row and  rodlike;  apical  portion  consists  of  a  broad,  flat  shaft,  about  the  same 
width  for  entire  length,  tapering  only  at  apex,  preapical  prominence  present 
on  the  ventral  edge,  obtuse-angled;  onl}-  slightly  curved;  apex  greatly  nar- 
rowed and  acutely  pointed,  very  lightly  chitinized;  strengthening  rod  ex- 
tends caudad  as  far  as  last  dorsal  tooth.  Toothed  area  on  dorsal  edge  occu- 
pies the  entire  length  of  the  apical  shaft ;  teeth  seventeen  to  eighteen  in  num- 
ber, medium  in  size,  somewhat  triangular  with  the  apex  flattened  and  the 
caudal  side  slightly  longer  than  the  cephalic,  regular  in  size  and  shape,  evenly 
spaced,  more  heavily  chitinized  than  the  rest  of  the  valve;  bear  a  large  num- 
ber of  small,  irregular,  secondary  teeth  on  both  edges;  tip  notched  with  small 
teeth  on  both  edges;  a  large  number  of  very  small  teeth  occur  between  the 
tip  of  the  preapical  prominence  and  the  extreme  apex;  ducts  rather  incon- 
spicuous, few  in  number  greatly  curved;  open  apically,  along  dorsal  edge  in 
and  near  teeth,  and  a  few  in  the  face  of  the  valve  back  from  the  edge. 

Cicadella  circellata  (Baker). 

(PI.  XXII,  fig.  8;   pi.  XXVI,  fig  8.) 

Length,  1.6  mm.;  greatest  width,  0.09  mm.  Apical,  flattened  portion  only 
slightly  wider  than  basal  portion;  preapical  prominence  wanting;  only  slightly 
curved,  tip  greatly  narrowed  but  rounded  at  extreme  apex,  chitinization 
rather  light  but  heavier  than  in  C.  hieroglyphica;  strengthening  rod  extends 
caudad  as  far  as  and  beyond  last  dorsal  tooth.    Toothed  area  on  dorsal  edge 


232  The  University  Science  Bulletin. 

occupies  nearly  all  of  broadened  area  or  about  half  the  total  length;  teeth 
thirteen  in  number,  rather  small,  in  the  general  shape  of  a  greatly  flattened 
obtuse  triangle  with  the  caudal  side  much  longer  than  the  cephalic,  regular 
in  spacing;  bear  secondary  teeth  on  both  edges,  five  to  eight  on  caudal  edge 
and  one  to  two  on  cephalic  edge,  those  on  the  caudal  edge  are  much  the 
larger;  tip  notched  with  small  teeth  on  both  dorsal  and  ventral  edges,  not  con- 
tinuous around  the  tip,  twenty  present  on  ventral  edge;  ducts  rather  incon- 
spicuous, though  more  conspicuous  than  in  C.  hieroglyphica,  elongate,  few  in 
number,  distinctly  curved;  open  apically,  along  dorsal  edge  in  and  near  teeth, 
and  in  face  of  valve  back  from  edge. 

Genus  Kolla  Distant. 

The  ovipositors  of  three  species  of  this  genus  have  been  examined 
and  found  to  be  similar.  In  each  case  the  ovipositor  consists  of  a 
curved,  narrow,  rodhke  basal  attachment  and  a  broader,  flat,  apical 
shaft  bearing  teeth  along  its  dorsal  edge.  The  teeth  are  triangular 
in  shape  and  bear  small  secondary  teeth  along  their  caudal  sides. 
The  tip  is  notched  in  each  case  with  small  teeth  present  on  both 
edges,  but  not  continuous  around  the  tip.  The  ducts  may  or  may 
not  be  visible;  when  visible  they  are  elongate,  rather  straight,  and 
few  in  number. 

Kolla  bifida  (Say). 

(PI.  XXII,  fig.   13;   pi.  XXVIII,  fig.   3.) 

Length,  1.5  mm.;  greatest  width,  0.22  mm.  Basal  connection  narrow  and 
rodlike ;  apical  shaft  broad,  flat,  narrowing  gradually  to  the  apex ;  no  preapical 
prominence ;  only  very  slightly  curved  beyond  curved  basal  attachment ;  apex 
narrowed  with  extreme  apex  rounded,  only  lightly  chitinized;  strengthening 
rod  extends  caudad  as  far  as  and  beyond  last  apical  tooth.  Toothed  area  on 
dorsal  edge  occupies  entire  length  of  apical  shaft  and  more  than  two-thirds  of 
the  entire  length;  teeth  fourteen  in  number,  medium  in  size,  in  the  general 
shape  of  a  greatly  flattened  obtuse  triangle  with  the  caudal  edge  much  longer 
than  the  cephalic  and  the  apex  rounded,  not  distinctly  more  heavily  chitinized 
than  the  rest  of  the  valve;  bear  small  secondary  teeth  along  the  caudal  edge 
only,  seven  to  twenty-one  on  each  tooth;  the  tip  is  notched  with  small  teeth 
on  both  dorsal  and  ventral  edges,  not  continuous  around  the  tip,  fourteen  to 
fifteen  on  ventral  edge;  ducts  inconspicuous,  only  apices  and  circular  open- 
ings visible;  open  apically,  along  the  doi-sal  edge  in  and  near  teeth,  and  a  few 
in  the  face  of  the  vahe  back  from  the  edge. 

Kolla  geometrica  (Signoret). 

(PI.  XXVIII,  fig.  2.) 

Length,  1.17  mm.;  greatest  width,  0.17  mm.  Basal  portion  narrow  and  rod- 
like; apical  shaft  broad,  flat,  tapers  to  apex;  no  preapical  prominence;  not 
curved  bej'ond  base;  tip  greatly  narrowed  with  extreme  apex  rounded  bluntly; 
chitinization  light  though  somewhat  heavier  than  in  K.  bifida;  strengthening 
rod  extends  caudad  as  far  as  last  dorsal  tooth.  Toothed  area  on  dorsal 
edge  occupies  entire  length  of  broadened  shaft;  teeth  seventeen  to  nineteen  in 


Readio:    Ovipositors  of  Cicadellid.e.  233 

number,  rather  small,  in  the  general  shape  of  a  greatly  flattened  obtuse  tri- 
angle with  the  caudal  side  much  longer  than  the  cephalic  and  the  apex  rounded, 
rather  irregular  in  size  and  spacing,  slightly  more  heavily  chitinized  than  the 
rest  of  the  valve;  bear  secondary  teeth  along  the  caudal  edge  only,  four  to 
fourteen  on  each  tooth;  tip  notched  with  teeth  on  both  dorsal  and  ventral 
edges,  not  continuous  around  the  tip,  thirteen  to  fifteen  on  ventral  edge;  ducts 
easily  visible  for  entire  length,  comparatively  few  in  number,  straight,  more 
numerous  apically;  open  apically,  along  dorsal  edge  in  and  near  teeth,  and  in 
the  face  of  valve  back  from  edge. 

•  Kolla  hartii  (Ball). 

(PI.  XXII,  fig.  14;   pi.  XXVIII,  fig.  1.) 

Length,  1.4  mm.;  greatest  width,  0.22  mm.  Basal  portion  narrow  and  rod- 
like; apical  shaft  broad  and  flat,  tapers  towards  apex;  no  preapical  promi- 
nence; not  curxed  beyond  base,  tip  greatly  narrowed,  extreme  apex  smoothly 
rounded,  lightly  chitinized;  strengthening  rod  extends  caudad  beyond  last 
doi-sal  tooth.  Toothed  area  on  dorsal  edge  occupies  entire  length  of  apical 
shaft;  teeth  seventeen  in  number,  medium  in  size,  in  the  shape  of  an  obtuse 
triangle,  though  not  so  flat  as  in  A',  bifida  and  A',  geometrica;  caudal  edge 
somewhat  longer  than  cephalic  edge,  rather  irregular  in  size  and  spacing,  not 
more  hea\'iiv  chitinized  than  the  rest  of  the  valve;  possess  small  secondary 
teeth  on  the  caudal  edge  of  each  primaiy  tooth,  eight  to  seventeen  to  each 
tooth;  tip  notched  with  small  teeth  on  both  dorsal  and  ventral  edges,  not  con- 
tinuous around  tip,  twenty-three  on  ventral  edge;  ducts  more  conspicuous  than 
in  K.  bifida,  less  conspicuous  than  in  A.  geometrica;  bases  and  apices  visible 
but  middle  portions  invisible;  open  apicalh',  along  dorsal  edge  in  and  near 
teeth,  and  in  face  of  valve  back  from  edge. 

Helochara  communis  Fitch. 

(PI.  XXII,  fig.  10;   pi.  XXVII,  fig.  5.) 

Length.  L44  mm.;  greatest  width,  0.22  mm.  Basal  portion  narrow  and 
ciu-ved,  apical  shaft  broad  and  flat,  about  the  same  width  for  entire  length, 
tapers  toward  tip;  bears  no  preapical  prominence;  not  curved  beyond  the 
base,  tip  greatlj^  narrowed,  with  the  extreme  apex  ending  in  an  obtuse  point, 
chitinization  mechimi;  strengthening  rod  extends  caudad  to  a  point  midway 
between  the  last  dorsal  tooth  and  the  tip.  Toothed  area  on  dorsal  edge  oc- 
cupies the  entire  length  of  the  apical  shaft,  nearly  the  entire  length  of  the 
valve;  teeth  thirty  in  number,  rather  large,  the  first  fourteen  of  a  general 
triangular  shape,  apical  sixteen  subquadrate  with  the  cephalic  side  longer  than 
the  caudal,  rather  irregular  in  size,  shape,  and  spacing;  basal  fourteen  primary 
teeth  bear  small  secondaiy  teeth  on  the  two  exposed  sides  of  the  triangle; 
apical  sixteen  primary  teeth  also  bear  secondary  teeth  on  the  outer  margin, 
three  to  fourteen  in  number;  tip  is  notched  with  small  teeth  on  both  doi^sal 
and  ventral  edges,  not  continuous  around  the  tip,  fort5'-six  present  on  venti-al 
edge;  ducts  conspicuoiLS,  rather  numerous,  elongate,  rather  straight,  occupy 
entire  shaft;  open  apically,  along  dorsal  edge  in  and  near  teeth,  and  in  face 
of  valve  back  from  edge. 


234  The  University  Science  Bulletin. 

■4 

Graphocephala  coccinca  (Forster). 

(PI.  XXII,  fig.  9;  pi.  XXVIII,  fig.  4.) 

Length,  1.87  mm.;  greatest  width,  0.22  mm.  Basal  portion  narrow  and  rod- 
hke,  apical  shaft  broad  and  flat,  about  the  same  width  for  entire  length;  pre- 
apical  prominence  present,  distinct,  obtuse-angled;  not  curved  bej-ond  base, 
tip  greatlj'  narrowed  and  ending  in  a  rather  blunt  point,  chitinization  light; 
strengthening  rod  e.xtends  caudad  beyond  last  dorsal  tooth.  Toothed  area  on 
dorsal  edge  occupies  entire  length  of  apical  shaft;  teeth  nineteen  in  number, 
medium  in  size,  of  a  general  triangular  shape  with  the  caudal  side  longer  than 
the  cephahc,  regular  in  size  and  shape,  distinctly  more  heavily  chitinized  than 
the  rest  of  the  valve;  bear  small  secondarj^  teeth  on  both  edges,  three  to  six 
on  cephalic  edge  and  eight  to  seventeen  on  caudal  edge;  tip  notched  with 
small  teeth  on  both  dorsal  and  ventral  edges,  not  continuous  around  the  tip, 
forty-five  between  preapical  prominence  and  extreme  apex;  ducts  easily  vis- 
ible though  not  conspicuous,  elongate,  rather  few  in  number,  curved,  most 
numerous  apically;  open  apically,  along  dorsal  edge  in  teeth,  one  to  two  to 
each  tooth,  and  a  few  in  the  face  of  valve  back  from  edge. 

Genus  Dr^culacephala  Ball. 

The  ovipositors  of  three  species  of  this  genus  have  been  examined 
and  found  to  be  similar.  The  ovipositor  of  Helochara  communis 
also  shows  its  close  relationship  to  this  genus.  In  each  case  the  ovi- 
positor consists  of  a  curved,  rodlike  basal  attachment  and  a  broad, 
fiat,  apical  shaft  which  bears  teeth  along  its  dorsal  edge  for  the  en- 
tire length.  The  distal  teeth  are  triangular  and  the  apical  teeth  sub- 
quadrate.  Both  types  of  teeth  bear  small  secondary  teeth.  The  tip 
in  every  case  is  notched  with  small  teeth  on  both  dorsal  and  ventral 
edges,  not  continuous  around  the  tip.  The  ducts  may  or  may  not 
be  conspicuous,  but  where  visible  are  elongate,  numerous  and  curved. 

Drceculacephala  mollipes  (Say). 

(PI.  XXII,  fig.  13  ;   pi.  XXVII,  fig.  2.) 

Length,  2.1  mm.;  greatest  width,  0.33  mm.  Basal  portion  narrow  and  rod- 
like, apical  shaft  broad  and  flat,  about  the  same  width  for  entire  length,  nar- 
rowing caudad  to  apex;  preapical  prominence  wanting;  not  curved  beyond 
base,  tip  greatly  narrowed  by  curving  ventral  edge  and  ends  in  an  obtuse  point, 
chitinization  moderately  heavy;  strengthening  rod  extends  caudad  to  a  point 
midway  between  last  dorsal  tooth  and  apex.  Toothed  area  on  dorsal  edge 
occupies  entire  length  on  apical  shaft;  teeth  twenty-eight  in  number,  large, 
basal  nine  of  a  general  triangular  shape,  apical  nineteen  subquadrate  with 
the  cephalic  side  longer  than  the  caudal,  rather  irregular  in  size,  shape  and 
spacing;  the  basal  nine  bear  small  secondary  teeth  on  both  cephalic  and 
caudal  edges;  the  apical  subquadrate  teeth  also  bear  secondary  teeth,  seven 
to  seventeen  on  the  outer  edge  and  all  except  the  apical  five  bear  secondary 
teeth  along  the  cephalic  edge,  one  to  six  in  number;  tip  is  notched  with  small 
teeth  on  both  dorsal  and  ventral  edges,  not  continuous  around  the  tip,  forty- 


Rkadio:    Ovipositors  of  Cicadellid.^.  235 

two  on  the  vent  ml  cnlge;  ducts  consijicuous,  rather  numerous,  elongate,  slightly 
curved,  most  numerous  apicalh-;  open  apically,  along  dorsal  edge  in  and  near 
teeth,  and  in  the  .•^in-face  of  the  valve  back  from  edge. 

Drceculaccyh  a  la  n  o  ve  boracensu  ( Fitch ) . 

(PI.  XXVII,  fig.  1.) 

Length,  2.65  mm.;  greatest  width,  0.33  mm.  Narrow  and  rodlike  at  base, 
broad  and  flat  ajiically,  apical  shaft  about  the  .same  width  for  entire  length; 
prcapical  prominence  wanting;  only  slightly  curved  beyond  base,  tip  greatly 
narrowed  by  slightly  curving  dorsal  edge  and  greatly  curving  ventral  edge 
and  ends  in  a  rather  sharp  point,  chitinization  light,  much  lighter  than  in  the 
other  members  of  this  genus  examined;  strengthening  rod  extends  cavidad  to 
a  point  midwaj'  between  last  dorsal  tooth  and  apex.  Toothed  area  on  dorsal 
edge  occupies  entire  length  of  apical  shaft,  the  dorsal  edge  bearing  the  teeth 
more  heavilj^  chitinized  than  the  rest  of  the  valve;  teeth  thirty-nine  to  forty- 
one  in  number,  of  medium  size,  the  basal  sixteen  of  a  general  triangular  shape, 
the  apical  twenty-three  subquadrate  wath  the  cephalic  side  longer  than  the 
caudal,  rather  irregular  in  size,  shape,  and  spacing;  the  basal  triangular  teeth 
bear  small  secondary  teeth  on  both  dorsal  and  ventral  edges;  the  apical  sub- 
quadrate  teeth  bear  three  to  twelve  secondary  teeth  on  the  outer  edge  and 
two  to  five  on  the  cephalic  edge;  the  secondary  teeth  midway  of  the  valve 
are  very  small  and  indistinct;  tip  notched  with  small  teeth  on  both  dorsal 
and  ventral  edges,  not  continuous  around  the  tip,  forty-five  present  on  the 
ventral  edge;  ducts  inconspicuous,  very  faintly  visible  except  at  openings, 
rather  numerous,  most  numerous  apically;  open  apically,  along  dorsal  edge 
in  and  near  teeth,  and  in  surface  of  valve  back  from  edge. 

Drceculacephala  reticulata  (Signoret). 

(PI.  XXII,  fig.  12;   pi.  XXVII,  fig.  3.) 

Length,  L75  mm.;  greatest  width,  0.27  mm.  Basal  portion  narrow  and  rod- 
like, apical  portion  broad,  fiat,  about  the  same  width  for  entire  length,  nar- 
rows at  apex;  preapical  prominence  wanting;  not  curved  beyond  curved  basal 
attachment,  tip  greatly  narrowed  by  slightly  .curved  dorsal  edge  and  greatly 
curved  ventral  edge,  ends  in  obtuse  point,  chitinization  medium,  less  heavy 
than  in  D.  noveboraccnsis  and  heavier  than  in  D.  mollipes;  strengthening  rod 
extends  caudad  to  a  point  midwaj^  between  last  dorsal  tooth  and  apex. 
Toothed  area  on  dorsal  edge  occupies  apical  shaft  for  entire  length;  teeth 
twenty-six  to  twenty-eight  in  number,  rather  large,  basal  eleven  of  a  general 
triangular  shape,  apical  fifteen  subquadrate  with  the  cephalic  side  longer 
than  the  caudal,  rather  irregular  in  size,  shape  and  spacing;  basal  eleven  bear 
small  secondai-y  teeth  on  both  edges;  apical  subquadrate  teeth  also  bear  sec- 
ondary teeth,  three  to  eleven  on  the  outer  edge  and,  with  the  exception  of 
the  apical  three,  one  to  five  on  the  cephalic  edge;  tip  notched  with  small 
teeth  on  both  dor.sal  and  ventral  edges,  not  continuous  around  the  tip,  42  pres- 
ent on  the  ventral  edge;  ducts  conspicuous,  rather  straight,  numerous,  most 
numerous  apically;  open  apically,  along  dorsal  edge  in  and  near  teeth,  and  a 
few  in  the  face  of  valve  back  from  edge. 


236  The  University  Science  Bulletin. 

Pagaronia  tripunctata  (Fitch). 

(PL  XXVII,  fig.   6.) 

Length,  1.5  mm.;  greatest  width.  0.17  mm.  Narrow  and  rodhke  at  base, 
apical  shaft  broad  and  flat,  about  the  same  width  for  entire  length,  narrows 
caudad  to  aj^ex;  preapical  prominence  wanting;  not  curved  beyond  base,  tip 
narrowed,  bkmtly  rounded  at  extreme  apex,  chitinization  Hght ;  strengthening 
rod  extends  caudad  as  far  as  last  dorsal  tooth.  Toothed  area  on  dorsal  edge 
occupies  entire  length  of  apical  shaft,  dorsal  edge  bearing  teeth  is  more  heavily 
chitinized  than  the  rest  of  the  valve;  teeth  seventeen  to  twenty  in  number, 
medium  in  size,  in  the  shape  of  a  flattened  obtuse  triangle  with  the  caudal 
side  much  longer  than  the  cephalic,  irregular  in  size,  shape  and  spacing;  bear 
secondary  teeth  on  both  outer  edges,  three  to  ten  on  cephalic  edge  and  three  to 
twenty-five  on  the  caudal  edge;  tip  notched  with  small  teeth  on  both  doreal 
and  ventral  edges,  not  continuous  around  the  tip,  30  present  on  ventral  edge; 
ducts  conspicuous,  rather  numerous,  only  slightly  curved,  most  numerous 
apically;  open  apically,  along  dorsal  edge  in  and  near  teeth,  and  a  few  in  the 
surface  of  valve  back  from  edge. 

Errhomenellu.s  montanus  Baker. 

(PI.  XXVII,  fig.   4.) 

Length,  2.83  mm.;  greatest  width,  0.22  mm.  About  the  same  width  for  en- 
tire length;  preapical  prominence  present,  rather  indistinct,  broadly  rounded; 
only  slightly  curved  beyond  curved  basal  attachment,  tip  greatly  narrowed 
with  the  extreme  apex  rounded,  chitinization  medium;  strengthening  rod  ex- 
tends caudad  as  far  as  fourth  dorsal  tooth.  Toothed  area  on  dorsal  edge  oc- 
cupies less  than  a  fourth  of  the  entire  length  apically;  teeth  seven  to  eight  in 
number,  small,  inconspicuous,  rounded,  regular  in  size  and  shape,  rather  far 
apart,  uneven  in  spacing,  bear  no  secondary  teeth;  tip  bears  no  teeth;  no  dis- 
tinct ducts  visible,  circular  openings  plainly  visible,  of  these  a  number  occur 
on  the  ventral  apical  edge,  some  at  the  extreme  apex,  and  also  along  the  en- 
tire dorsal  edge. 

Subfamily  GYPONINtE   (Stal). 
Genus  Gypona  Germar. 

The  ovipositors  of  four  species  of  this  genus  have  been  examined 
and  found  to  be  similar.  The  ovipositor  of  Xerophloea  liridis,  the 
only  other  species  of  this  subfamily  examined,  is  not  similar  to  the 
ovipositors  of  the  genus  Gypona.  In  this  genus  the  ovipositor  is 
relatively  short,  stout  and  broad,  usually  being  rather  heavily  chi- 
tinized. The  apical  portion  is  not  much,  if  any,  wider  than  the  basal 
portion.  Teeth  are  present  along  the  dorsal  edge  for  from  one- 
fourth  to  one-half  the  apical  length.  The  tip  may  or  may  not  bear 
teeth  on  one  or  both  edges.  The  ducts  in  every  case  are  conspicuous, 
large,  elongate  and  numerous. 


Readio:    Ovipositors  of  Cicadellid.^.  237 

Gypona  octo-lineata  (Say). 

(PI.  XXIII,  fig.   1;   pi.  XXVIII,  fig.  8.) 

Length,  2  iiun.;  greatest  width,  0.48  mm.  Consists  of  a  rather  narrow  basal 
portion,  which  widens  out  into  a  broad,  apical  portion,  which  narrows  caudad 
to  the  apex,  point  of  greatest  width  about  midway;  no  preapical  prominence; 
shghtiy  curved,  tip  greatly  narrowed  with  the  extreme  apex  broadly  rounded, 
chitinization  moderately  heavy;  strengthening  rod  extends  caudad  almast  to 
apex.  Toothed  area  on  dorsal  edge  occupies  about  a  half  of  the  apical  length; 
teeth  numerous,  rather  small,  rounded,  irregular  in  size,  shape-  and  spacing; 
bear  no  secondary  teeth;  tip  bears  small  teeth  on  both  dorsal  and  ventral 
edges,  not  continuous  around  the  tip,  thirteen  on  ventral  edge;  ducts  con- 
spicuous, large,  numerous,  apparently  arising  from  a  common  cavity  in  the 
center  of  the  valve,  most  numerous  at  the  tip;  open  along  ventral  apical  edge, 
at  extreme  apex,  and  along  the  dorsal  edge  for  the  entire  length,  though  very 
few  in  basal  region. 

Gypotia  biniaculata  Spangberg. 

(PI.  XXVIII,  fig.  5.) 

Length,  3.17  mm.;  greatest  width,  0.42  mm.  About  the  same  width  for  en- 
tire length,  bears  a  broadly  rounded,  heavily  chitinized  prominence  on  the  dor- 
sal edge  somewhat  less  than  midway  of  the  length,  tapers  caudad  toward  apex; 
slightly  curved,  tip  gradually  naiTOwed,  ending  in  a  blunt  point,  chitinization 
hea\-y,  much  hea\  ier  than  in  G.  octo-lineata;  strengthening  rod  extends  caudad 
almost  to  apex.  Toothed  area  on  dorsal  edge  occupies  about  one-third  of 
apical  length;  teeth  numerous,  rather  small,  rounding,  iiTegular  in  size  and 
shai)e;  bear  no  secondaiy  teeth;  tip  notched  on  dorsal  edge  and  at  extreme 
apex  with  small  teeth,  none  present  on  ventral  edge;  ducts  conspicuous,  very 
large,  rather  straight;  open  apically  and  along  dorsal  and  ventral  edges  for 
the  length  of  the  toothed  area,  a  few  also  open  in  the  basal  region. 

Gypona  angulata  Spangberg. 

(PI.  XXVIII,  fig.  6.) 

Length,  L75  mm.;  greatest  width,  0.35  mm.  Apical  portion  somewhat  wider 
than  basal,  point  of  greatest  width  about  midway,  tapers  caudad  to  apex; 
only  slightly  curved,  tip  gradually  narrowed  with  the  extreme  apex  rounded, 
chitinization  moderately  heavy;  strengthening  rod  extends  caudad  to  within 
a  short  distance  of  tip.  Toothed  area  extends  from  the  prominence  on  the 
dorsal  edge  to  the  apex;  teeth  numerous,  small  and  indistinct,  rather  flatly 
rounded,  verj'  irregular  in  size,  shape  and  spacing;  bear  no  secondary  teeth; 
no  teeth  on  tip;  ducts  consi;Hcuous,  large,  numerous,  broadly  curving,  most 
numerous  apicall}^;  open  apically,  along  ventral  apical  edge,  along  dorsal 
edge  of  toothed  area,  and  a  few  in  the  basal  portion  of  the  valve. 

Gypona  pectoralis  Spangberg. 

(PI.  XXVIII,  fig.  7.) 

,  Length,  L83  mm.;  greatest  width,  0.42  mm.  Very  broad  and  heavy  for 
entire  length,  point  of  greatest  width  about  midway,  where  there  is  a  broadly 
rounded  prominence  on  the  dorsal  edge,  narrows  caudad  to  apex;  slightly 
curved,  tip  greatly  narrowed  with  extreme  apex  rounded,  chitinization  very 


238  The  University  Science  Bulletin. 

heavy;  strengthening  rod  extends  caudad  almost  to  apex.  Toothed  area  on 
dorsal  edge  occupies  about  a  fourth,  of  the  apical  length ;  teeth  very  few  in 
number,  small,  flat,  somewhat  rounded,  very  irregular  in  size,  shape  and 
spacing,  bear  no  secondary  teeth;  no  teeth  present  at  extreme  apex;  ducts 
conspicuous,  large,  somewhat  branching,  more  numerous  apically;  open 
apically,  along  ventral  apical  edge,  and  along  dorsal  for  entire  length. 

Xerophloea  viridis  (Fabricius). 

(PL  XXIII,  fig.  2;  PI.  XXIX,  fig.  2.) 

Length,  2.88  mm.;  greatest  width,  0.08  mm.  Narrow  and  rodlike,  about  the 
same  width  for  entire  length;  distinctly  curved,  tip  bluntly  rounded,  chitiniza- 
tion  medium;  strengthening  rod  extends  caudad  as  far  as  twentieth  dorsal  tooth. 
Toothed  area  on  dorsal  edge  occupies  a  little  less  than  the  apical  third;  teeth 
twenty-four  in  number,  medium  in  size,  rounded,  regular  in  size  and  shape, 
the  apical  four  are  farther  apart  than  the  basal  twenty;  the  apical  three  bear 
one  to  three  secondary  teeth  on  the  caudal  edge;  tip  notched  with  small  teeth 
on  both  dorsal  and  ventral  edge,  not  continuous  around  the  tip,  twelve  to 
thirteen  on  ventral  edge;  ducts  conspicuous,  relatively  few  in  number,  most 
numerous  at  apex;  open  apically  and  along  entire  dorsal  edge,  even  in  basal 
portion. 

Subfamily  JASSIN./E  (Amyot  and  Serville). 

Tribe  Acucephalini  (Dohrn). 

Stroggylocephalus  agrestis   (Fallen). 

(PI.   XXIX,   fig.    3.) 

Length,  L88  mm.;  greatest  width,  0.17  mm.  Curved  base  rather  narrow 
and  rodlike,  rather  broad  and  flat  beyond  base,  about  the  same  width  for 
entire  length,  tapers  gradually  toward  apex;  curved  at  base  of  broadened  area, 
tip  greatly  narrowed  with  extreme  apex  narrowly  rounded,  chitinization  rather 
light,  strengthening  rod  extends  caudad  as  far  as  sixth  dorsal  tooth  from  apex. 
Toothed  area  on  dorsal  edge  occupies  a  little  less  than  half  the  entire  length, 
teeth  twenty-six  to  twenty-eight  in  number,  rather  small,  of  a  general  tri- 
angular shape,  some  roimded  and  some  sharply  pointed,  verj'  irregular  in  size, 
shape  and  spacing;  bear  no  secondary  teeth;  tip  notched  with  a  few  small, 
indistinct,  irregular  teeth  on  the  ventral  edge  only;  ducts  conspicuous,  elongate ; 
open  along  ventral  apical  edge,  at  extreme  apex,  and  along  entire  dorsal  edge; 
the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a  distinct,  elongate, 
heavily  chitinized  connection  present  on  the  dorsal  edge  near  the  base  of  the 
broadened  area. 

Memnonia  consobrina  Ball. 

(PI.   XXIX,   fig.   5.) 

Length,  2.2  mm.;  greatest  width,  0.25  mm.  Gunned  basal  attachment  nar- 
row and  rodlike,  beyond  base  is  a  rather  narrow,  lightly  chitinized  area  ex- 
tending caudad  for  about  a  third  of  the  length;  caudad  of  this  is  a  broader, 
more  heavily  chitinized  portion  occupying  the  rest  of  the  valve;  narrows, 
rather  abruptly  caudad  to  apex;  only  slightly  curved  beyond  base,  tip  greatly 
narrowed  by  broadly  cui-ving  doi-sal  edge;  extreme  apex  ends  in  sharp,  acute- 


Readio:    Ovipositors  of  Cicadellid.e.  239 

angled  point,  rhitiuization  nioderatt'ly  light;  strengthening  rod  extends  caudad 
almost  to  apex.  There  are  no  evidences  of  distinct  teeth,  though  the  margin 
is  irregularly  roughened  in  se\eral  i)laces;  area  of  ducts  conspicuous,  granular 
in  appearance,  circular  duct  ojienings  visible;  ope>n  along  ventral  apical  edge, 
at  extreme  apex,  and  along  dorsal  edge  for  entire  length. 

Xestocephahis  pnlicarius  Van  Duzee. 

(PI.  XXIII,  fig.  3;    pi.  XXIX,  fig.   1.) 

Length.  0.88  mm.;  greatest  width,  0.08  mm.  Curved  basal  attachment 
narrow  and  rodlike,  rest  of  valve  about  the  same  width  for  entire  length; 
preapical  prominence  on  ventral  edge  present,  distinct,  obtuse  angled,  giv- 
ing the  apical  portion  of  the  valve  the  appearance  of  a  spear  head;  tip 
narrowed  abruptly  beyond  preapical  prominence;  very  distinctly,  broadly 
and  evenly  curved;  tip  greatly  narrowed,  ending  in  sharply  pointed  ex- 
treme apex,  chitinization  medium ;  strengthening  rod  extends  caudad  to 
within  a  short  distance  of  extreme  apex;  dorsal  edge  of  tip  bears  an  indistinct 
notch  where  rod  meets  dorsal  edge.  Toothed  area  on  dorsal  edge  occupies 
the  apical  three-fourtlxs  of  the  length;  teeth  fourteen  to  fifteen  in  number, 
small,  indistinct,  rather  irregular  in  size,  shape  and  spacing,  especially  basally; 
bear  no  secondary  teeth;  tip  bears  no  teeth;  ducts  rather  inconspicuous  though 
plainly  visible,  rather  few;  open  in  preapical  prominence,  at  extreme  apex, 
and  along  entire  dorsal  edge;  the  two  valves  of  the  pair  are  joined  one  to 
the  other  by  a  distinct,  elongate,  narrow,  heavily  chitinized  connection  present 
on  the  dorsal  edge,  this  connection  more  liberally  supplied  with  ducts  than  the 
portion  of  the  valve  immediately  caudad  of  it. 

Tribe  Jassini  (Dohrn). 
Dorycephalus  platyrhynchus  Osborn. 

(PI.   XXIX,   fig.   4.) 

Length,  3.5  mm.;  greatest  width,  0.5  mm.  Curved  basal  portion  narrow 
and  rodlike ;  at  a  point  about  a  third  of  the  length  it  widens  into  a  broad, 
flat  shaft  which  tajiers  slightly  caudad;  point  of  greatest  width  is  a  little  less 
then  midway;  only  slightly  cui-ved  beyond  base,  tip  greatly  narrowed  by 
rounding  dorsal  edge,  extreme  apex  rounded,  chitinization  very  light;  strength- 
ening rod  extends  caudad  to  within  a  short  distance  of  the  extreme  apex. 
There  are  no  distinct  teeth,  though  the  ventral  apical  edge  and  the  entire 
dorsal  edge  are  slightly  and  irregularly  roughened;  area  of  ducts  inconspicu- 
ous, no  ducts  visible,  circular  duct  openings  visible;  a  single  row  opens  along 
the  ventral  apical  edge,  a  single  row  along  the  dorsal  apical  edge,  distad 
of  these  a  double  row,  distad  of  the  double  row  a  triple  row,  and  still  further 
distad  in  the  wider  portion  of  the  valve  as  many  as  four  or  five  irregular  rows 
open  in  the  entire  breadth  of  the  valve. 

Hecalus  lineatus  (Uhler). 

(PI.   XXIX,  fig.  8.) 

Lenth,  4.1  mm.;  greatest  width,  0.43  mm.  Curved  basal  portion  narrow  and 
rodlike,  widens  bej'ond  base  into  broa<l,  flat  portion  which  tapers  gradually 
caudad  to  apex,  point  of  greatest  width  about  two-fifths  of  length;  only  slightly 
curved  bej'ond  base,  tip  greatly  narrowed,  ending  in  sharp  point    chitinization 


240  The  University  Science  Bulletin, 

moderately  light;  strengthening  rod  extends  caudad  to  within  a  short  distance 
of  the  apex.  No  distinct  teeth  present,  the  ventral  apical  edge  and  the  entii-e 
dorsal  edge  except  at  the  apex  are  slightly  and  irregularly  roughened;  area  of 
ducts  conspicuous,  granular  in  appearance,  circular  duct  openings  visible;  open 
along  ventral  apical  edge  and  along  dorsal  edge  for  entire  length,  in  the  basal 
region  they  open  irregularly  in  the  entire  width  of  the  valve. 

Sipanghergiella  mexicana  Baker. 

(PI.  XXIX,  fig.   6.) 

Length,  1.9  mm.,  greatest  width,  0.27  mm.  Cvu-ved  base  narrow  and  rodlike, 
widens  into  broad,  flat  portion,  about  the  same  width  for  entire  length,  nar- 
rowing caudad  to  apex;  bears  two  prominences,  one  on  dorsal  edge  somewhat 
less  than  midway,  and  the  other  on  the  ventral  edge  about  two-thirds  the 
length,  more  angular;  not  curved  beyond  the  base,  narrowed  at  tip  evenly  and 
gradually  by  both  curving  edges,  ends  in  sharp  point,  chitinization  medium; 
strengthening  rod  extends  caudad  almost  to  apex.  Bears  no  teeth;  area  of 
ducts  granular  in  appearance,  circular  openings  very  small  and  inconspicuous; 
open  along  ventral  apical  edge,  at  extreme  apex,  along  dorsal  edge,  and  in  basal 
portion  of  valve. 

Parabolocratus  ^avidus  Signoret. 

(PI.  XXIX,  fig.   7.) 

Length,  2.3  mm.;  greatest  width,  027  mm.  Curved  base  narrow  and  rodlike, 
widens  gradually  into  broad,  flat  portion,  about  the  same  width  for  entire 
length,  narrowing  only  at  apex;  bears  two  broadly  rounded  prominences,  one 
on  the  dorsal  edge  about  a  third  of  the  length,  the  other  on  the  ventral  edge 
about  two-thirds  the  length;  only  slightly  curved  beyond  base,  tip  gradually 
narrowed  and  ends  in  sharp  point,  chitinization  medium;  strengthening  rod 
extends  caudad  to  within  a  short  distance  of  the  apex.  Toothed  only  at  ex- 
treme tip;  teeth  are  continuous  around  the  tip,  of  these  six  are  on  the  dorsal 
edge  and  ten  on  the  ventral  edge;  ducts  inconspicuous,  invisible  except  at  ex- 
treme apices,  circular  openings  visible ;  open  along  ventral  apical  edge  and 
along  entire  dorsal  edge  except  in  dorsal  prominence. 

Aligia  jucunda   (Uhler). 

(PI.  XXIX,  fig.   11.) 

Length,  L83  mm.;  greatest  width,  0.17  mm.  Curved  base  narrow  and  rod- 
like, slightly  wider  beyond  base  for  a  little  more  than  half  the  length;  apical 
portion  of  valve  widens  abruptly,  tapers  caudad  to  apex;  distinctly  curved,  tip 
narrowed  by  curved  dorsal  edge,  extreme  apex  bluntly  rounded,  almost  square, 
chitinization  medium;  strengthening  rod  extends  caudad  to  within  fourteen 
dorsal  teeth  of  apex.  Toothed  area  on  dorsal  edge  occuj^ies  somewhat  less  than 
the  apical  half;  teeth  eighty-eight  in  number,  very  small,  wedge-shaped,  fairly 
regular  in  size,  shape  and  spacing,  bear  small  secondaiy  teeth  on  outer  edge, 
one  to  four  in  number;  tip  devoid  of  teeth  on  ventral  and  caudal  edges;  area 
of  ducts  conspicuous,  granular  in  appearance,  though  a  few  elongate  ducts  are 
visible  apically;  ducts  straight,  numerous;  open  along  ventral  apical  edge,  at 
extreme  apex,  along  dorsal  edge  of  toothed  area,  and  irregularly  in  the  basal 
region.    The  two  valves  of  the  pair  are  joined  one  to  the  other  by  an  elongate 


Readio:    Ovipositors  of  Cicadellid.e.  241 

connection  only  slightly  more  heavily  chitinized  than  the  rest  of  the  valve, 
present  on  the  dorsal  edge  in  the  basal  region. 

Genus  Mesamia  Ball. 

Two  species  of  this  genus  have  been  examined  and  found  to  be 
similar.  In  each  case  the  ovipositor  consists  of  a  curved  basal  at- 
tachment, a  somewhat  wider  portion  that  extends  about  to  the  mid- 
point, and  an  apical  portion  which  bears  teeth  along  its  dorsal  edge. 
A  small  preapical  prominence  is  present  on  one  of  the  two  species. 
Some  of  the  primary  teeth,  at  least,  bear  secondary  teeth.  In  one 
species  the  tip  bears  distinct  teeth,  in  the  other  the  tip  is  only  irreg- 
ularly roughened.  The  area  of  ducts  is  granular  in  appearance  and 
the  duct  openings  are  visible.  The  two  valves  of  the  pair  are  joined 
one  to  the  other  by  an  elongate,  heavily  chitinized  connection  pres- 
ent on  the  dorsal  edge  in  the  basal  region. 

Mesamia  straminea  (Osborn). 

(PI.  XXIX,  fig.   9.) 

Length,  1.53  mm.;  greatest  width,  0.18  mm.  Curved  basal  portion  narrow 
and  rodlike,  bej^ond  base  is  a  wider,  heavity  chitinized  area  extending  almost  to 
the  midpoint;  the  apical  toothed  area  occupies  the  rest  of  the  valve  and  is 
slightly  wider,  less  heavily  chitinized,  and  tapers  caudad  to  the  apex;  bears  no 
preapical  prominence;  slightly,  though  distinctly  curved,  tip  narrowly  rounded, 
chitinization  moderately  heavy;  strengthening  rod  extends  caudad  as  far  as 
fourth  dorsal  tooth  from  the  tip.  Toothed  area  on  dorsal  edge  occupies  the 
apical  half;  about  twenty-five  primaiy  teeth  present,  of  medium  size,  very 
irregular  as  to  size,  shape  and  spacing;  the  apical  ten  are  somewhat  regular, 
long  and  flatly  rounded  and  may  or  ma}^  not  bear  a  few  secondaiy  teeth  on  the 
caudal  edge,  the  teeth  distad  of  these  extremely  irregular;  the  tip  is  irregularly 
roughened  on  both  dorsal  and  ventral  edges,  but  bears  no  distinct  teeth;  area 
of  ducts  granular  in  appearance,  duct  openings  easily  visible;  open  along  ven- 
tral apical  edge,  at  extreme  apex,  along  dorsal  edge  of  toothed  area,  and  in  the 
basal  portion  of  the  valve;  the  two  valves  of  the  pair  are  joined  one  to  the 
other  by  a  distinct,  elongate,  heavily  chitinized  connection  present  on  the 
dorsal  edge  at  about  one-third  the  length;  the  apex  is  apparently  strengthened 
by  a  straight,  narrow  rod  that  extends  from  the  interior  of  the  valve  to  the 
extreme  apex,  occupying  about  the  apical  .sixth  of  the  length. 

Mesamia  vitellina  (Fitch). 

rPI.   XXIII,  fig.   7;    pi.  XXIX,   fie.    10.) 

Length,  1.47  mm.;  greatest  width,  0.13  mm.  Curved  basal  connection  nar- 
row and  rodlike,  widens  out  into  broader  area,  which  is  about  the  same  width 
for  entire  length  and  toothed  apically;  a  small  preapical  prominence  is  present 
on  the  ventral  edge;  distinctlj^  curved,  tip  narrowed,  narrowly  rounded  at  ex- 
treme apex,  chitinization  moderately  heavy;  strengthening  rod  extends  caudad 
as  far  as  fifth  dorsal  tooth  from  apex.  Toothed  area  on  dorsal  edge  occupies 
a  little  less  than  the  apical  half;  teeth  seventeen  in  number,  rather  small,  in 


242  The  Univeksity  Science  Bulletin. 

the  general  shape  of  a  greatlj^  flattened  obtuse  triangle  with  a  rounded  apex, 
fairly  regular  in  size,  shape  and  spacing,  those  distad  being  very  flat  and  in- 
distinct, bear  small;  numerous  secondary  teeth  on  both  cephalic  and  caudal 
edges;  tip  notched  with  small  teeth  on  both  edges,  not  continuous  around  the 
tip,  twelve  to  thirteen  present  between  preapical  prominence  and  apex,  area 
of  ducts  granular  in  appearance,  circular  duct  openings  visible;  open  along 
ventral  apical  edge,  at  extreme  apex,  along  dorsal  edge  of  toothed  area,  and 
in  basal  portion  of  the  valve;  the  two  valves  of  the  pair  are  joined  one  to  the 
other  by  an  elongate,  distinct,  heavily  chitinized  connection  present  on  the 
dorsal  edge  at  a  point  a  little  more  than  one-third  the  length  of  the  valve ;  the 
apex  is  apparently  strengthened  by  a  short,  narrow  rod  which  can  be  seen 
only  at  the  extreme  apex,  much  shorter  and  lighter  than  in  M.  straminca. 

Genus  Scaphoideus  Uhler. 

The  ovipositors  of  two  species  of  this  genus  have  been  examined. 
They  do  not  resemble  each  other  to  the  degree  that  is  common  be- 
tween two  species  of  the  same  genus,  though  they  have  many  points 
in  common.  In  each  case  the  ovipositor  is  about  the  same  width  for 
the  entire  length,  narrows  only  at  the  apex,  and  bears  no  preapical 
prominence.  The  dorsal  edge  bears  teeth  along  the  apical  two-fifths 
of  its  length,  but  these  teeth  differ  in  number,  size,  shape,  and  pos- 
session of  secondary  teeth  in  the  two  species,  the  tip  bears  teeth  in 
one  species,  none  in  the  other;  the  area  of  ducts  is  granular  in  ap- 
pearance and  the  duct  openings  are  visible.  The  two  valves  of  the 
pair  are  joined  one  to  the  other  by  an  elongate,  heavily  chitinized 
connection  present  on  the  dorsal  edge  about  midway  of  the  valve. 

Scaphoideus  scalaris  Van  Duzee. 

(PL   XXX,   fig.    1.) 

Length,  1.53  mm.;  greatest  width,  0.13  mm.  Curved  basal  connection  nar- 
row and  rodlike,  rest  of  valve  about  the  same  width  for  entire  length,  narrow- 
ing only  at  apex;  beai*s  no  preapical  prominence;  slightly  curved,  tip  narrowed, 
bluntly  rounded  at  extreme  apex,  chitinization  rather  heavy ;  strengthening  rod 
extends  caudad  almost  to  second  dorsal  tooth  from  apex.  Toothed  area  on 
dorsal  edge  occupies  a  little  more  than  the  apical  two-fifths  of  the  length; 
teeth  nine  in  number,  rather  large,  broadlj^  rounded,  rather  irregular  in  size, 
shape  and  spacing  being  farther  apart  basally;  bear  no  secondary  teeth;  tip 
bears  no  teeth,  ducts  granular  in  appearance,  circular  duct  openings  visible; 
open  along  ventral  apical  edge,  at  extreme  apex,  along  dorsal  edge  of  toothed 
area,  and  irregularly  in  the  basal  region  where  elongate  ducts  are  visible;  the 
two  valves  of  the  pair  are  joined  one  to  the  other  by  an  elongate,  heavily 
chitinized  connection  present  on  the  dorsal  edge  about  midway  of  the  valve ; 
this  connection  occupies  a  little  less  than  a  third  of  the  entire  length  and  bears 
the  first  dorsal  tooth. 


Readio:    Ovipositors  of  Cicadellid.e.  248 

Scaphoideiis  immistus  (Say). 

(PI.  XXIII,  fig.  4;  pi.  XXX,  fig.  2.) 

Length,  1.77  mm.;  greatest  width,  0.11  mm.  Narrow  and  rodliko  at  base, 
rest  of  vahe  somewhat  broader,  about  the  same  width  for  entire  length,  nar- 
rows caudad  to  ape.\;  bears  no  preapical  prominence;  distinctly  curved,  tip 
greatly  narrowed,  extreme  apex  rounded,  chitinization  moderately  heavy; 
strengthening  rod  extends  caudad  as  far  as  fifth  dorsal  tooth  from  apex. 
Toothed  area  on  dorsal  edge  occupies  a  little  less  than  the  apical  two-fifths 
of  the  length;  teeth  sixteen  in  number,  rather  small,  rounded,  rather  irregular 
in  size,  shape  and  spacing  being  farther  apart  basally;  may  bear  secondary  on 
one  or  both  edges,  small  secondarj-  teeth  also  pi-esent  on  margin  of  valve  be- 
tween primary  teeth,  general  arrangement  of  secondary  teeth  very  irregular; 
tip  notched  with  small,  irregular  teeth  on  both  edges,  though  more  distinct  on 
the  ventral  edge,  not  continuous  around  the  tip,  eight  to  nine  present  on  the 
ventral  edge;  ducts  partly  granular  and  pai'tly  elongate,  circular  openings  visi- 
ble ;  open  along  ventral  apical  edge,  at  extreme  apex,  along  dorsal  edge  of 
toothed  area,  and  irregularly  in  the  basal  region;  the  two  valves  of  the  pair 
are  joined  one  to  the  other  by  an  elongate,  heavily  chitinized  connection 
present  on  the  dorsal  edge  about  midway,  occupies  about  two-fifths  of  length, 
bears  one  rounded  tooth  somewhat  caudad  of  the  center. 

Genus  Platymetopius  Burmeister. 

The  ovipositors  of  three  members  of  this  genus  have  been  ex- 
amined and  found  to  be  similar.  In  each  case  the  basal  half  is 
slightly  narrower  than  the  apical  half,  which  bears  teeth  along  its 
dorsal  edge.  A  preapical  prominence  may  or  may  not  be  present. 
The  primary  teeth  bear  secondary  teeth  on  both  edges,  more  on  the 
caudal  than  on  the  cephalic  edge.  In  every  case  the  tip  is  notched 
with  small  teeth  on  both  dorsal  and  ventral  edges;  these  may  or 
may  not  be  continuous  around  the  tip.  The  area  of  ducts  may  or 
may  not  be  conspicuous,  when  visible  is  granular  in  appearance  with 
the  duct  openings  visible.  The  two  valves  of  the  pair  are  joined 
one  to  the  other  by  a  rather  short,  chitinous  connection  present  on 
the  dorsal  edge  of  the  basal  area. 

Platy77ietopius  acutus  (Say). 

(Pl.  XXIII,  fig.  5;    pi.  XXX,  fig.  3.) 

Length,  1.36  mm.;  greatest  width,  0.17  mm.  Curved  basal  connection  nar- 
row and  rodlike;  beyond  this  is  a  somewhat  wider  portion  extending  nearly  to 
midpoint,  and  apically  the  toothed  area,  about  the  same  width  for  entire 
length,  narrowing  only  at  apex,  bears  an  indistinct,  broadly  angled  preapical 
prominence  on  the  ventral  edge;  slightly  though  distinctly  curved,  tip  nar- 
rowed and  ending  in  a  bluntly  rounded  extreme  apex,  chitinization  light; 
strengthening  rod  extends  caudad  as  far  as  fifth  dorsal  tooth  from  apex. 
Toothed  area  on  dorsal  edge  occupies  the  apical  half  of  the  valve;  teeth 
twenty-seven  in  number,  rather  large;  in  the  greater  number  of  teeth  the  basal 


244  The  University  Science  Bulletin. 

edges  of  each  tooth  are  more  or  less  parallel  and  the  apical  edges  taper,  giving 
a  gabled  appearance  to  the  tooth;  the  caudal  edge  is  somewhat  longer  than 
the  cephalic,  fairly  regular  as  to  size,  shape  and  spacing,  though  not  entirely  so; 
primary  teeth  bear  small  secondary  teeth  on  caudal  edge  more  commonly, 
and  a  few  teeth  also  bear  secondary  teeth  on  the  cephalic  edge;  tip  notched 
with  small  teeth  on  both  dorsal  and  ventral  edges,  continuous  around  the  tip, 
nineteen  present  on  ventral  edge  between  preapical  prominence  and  apex; 
area  of  ducts  inconspicuous,  the  apices  and  circular  openings  alone  being 
visible;  open  along  ventral  ai)ical  edge,  at  extreme  apex,  along  dorsal  edge  of 
toothed  area,  and  irregularly  in  the  basal  portion  of  the  valve;  the  two  valves 
of  the  pair  are  joined  one  to  the  other  by  an  indistinct,  chitinous  connection 
present  on  the  dorsal  edge  at  about  a  third  of  the  length. 

Platymetopius  cinereus  Osborn. 

(PI.    XXX,    fig.    4.) 

Length,  1.13  mm.;  greatest  width,  0.13  mm.  Curved  basal  connection  nar- 
row and  rodlike;  a  wider  area  extends  almost  to  midpoint,  a  slighter  wider 
apical  portion  occupies  the  rest  of  the  valve;  no  preapical  prominence; 
slightly  curved,  tip  narrowed,  ends  in  obtuse-angled  point,  chitinization  mod- 
erately light;  strengthening  rod  extends  caudad  as  far  as  sixth  dorsal  tooth 
from  apex.  Toothed  area  on  dorsal  edge  occupies  a  little  more  than  the  apical 
half;  teeth  twenty-three  to  twenty-four  in  number,  moderately  large,  of  a 
general  triangular  shape  with  the  caudal  side  in  most  cases  longer  than  the 
cephalic  and  with  apices  rounded,  fairly  regular  in  size,  shape  and  spacing; 
bears  small  secondary  teeth  on  caudal  edge,  three  to  eight  in  number,  and  a  few 
of  the  teeth  also  bear  a  single  secondary  tooth  on  the  cephalic  edge;  tip 
notched  with  small  teeth  on  both  dorsal  and  ventral  edges,  more  numerous  and 
distinct  on  ventral  edge,  not  continuous  around  the  tip,  fourteen  present  on 
the  ventral  edge;  ducts  very  inconspicuous,  only  a  few  being  visible  in  the 
apex  of  the  valve,  apices  of  ducts  and  duct  openings  visible ;  open  along  ventral 
apical  edge,  at  extreme  apex,  along  dorsal  edge  of  toothed  area,  and  irregu- 
larly in  the  basal  portion  of  the  valve;  the  two  valves  of  the  pair  are  joined 
one  to  the  other  by  a  poorly  defined,  heavily  chitinized  connection  present  on 
the  dorsal  edge  at  about  one-third  the  length. 

Platyjnetopius  frontalis  Van  Duzee. 

(PI.  XXIII,  fig.  6;   pi.  XXX,  fig.  5.) 

Length,  1.26  mm.;  greatest  width,  0.16  mm.  Curved  basal  connection  nar- 
row and  rodlike;  a  slightly  wider  portion  extends  to  a  point  nearly  midway; 
the  apical  portion  is  slightly  wider,  naiTOws  caiidad  to  apex,  bears  a  suggestion 
of  a  broadly  rounded  preapical  prominence  on  the  ventral  edge;  shghtly 
curved,  tip  narrowed,  ends  in  a  blunt,  obtuse-angled  point,  chitinization  mod- 
erately heavy,  heavier  than  in  P.  acutus  and  P.  cinereus;  strengthening  rod 
extends  caudad  as  far  as  fifth  dorsal  tooth  from  apex.  Toothed  area  on  dorsal 
edge  occupies  a  little  more  than  the  apical  half;  teeth  twenty-one  to  twenty- 
two  in  number,  rather  large;  in  the  greater  number  the  basal  edges  of  each 
tooth  are  parallel  and  the  apical  edges  taper  to  a  rounded  apex,  fairly  regular 
in  size,  shape  and  spacing ;  bear  small  secondary  teeth  for  the  most  part  on  the 


Readio:    Ovipositors  of  Cicadellid.e.  245 

caudal  edge;  tip  iiotrliod  with  tiiuall  teeth  ou  both  dorsal  and  ventral  edges, 
practically  continuous  around  the  tip,  eighteen  present  on  the  ventral  edge; 
area  of  ducts  conspicuous,  granular  in  appearance,  circular  duct  openings  vis- 
ible: open  along  ventral  apical  edge,  along  dorsal  edge  of  toothed  area,  and 
irregularly  in  the  basal  area  of  the  valve;  the  two  valves  of  the  pair  are  joined 
one  to  the  other  by  a  distinct,  rectangular,  heavily  chitinized  connection  pres- 
ent on  the  doi-sal  edge  at  about  one-third  the  length. 

Genus  Deltocephalus  Burmeister. 
The  u\ipositors  of  seven  species  of  this  genus  have  been  examined 
and  found  to  be  similar.  The  ovipositor  is  slightly  curved,  gradually 
narrowed  toward  the  apex,  with  apical  half  little  if  any  wider  than 
the  basal  half.  The  apical  half  bears  teeth  along  its  dorsal  edge, 
usually  small,  rounded,  and  bearing  a  few  secondary  teeth.  The 
apex  may  or  may  not  bear  small  teeth  on  one  or  both  edges.  The 
area  of  ducts  may  or  may  not  be  conspicuous ;  when  visible  is  gran- 
ular in  appearance  with  duct  openings  visible.  The  two  valves  of 
the  pair  are  joined  one  to  the  other  by  a  chitinous  connection  pres- 
ent on  the  dorsal  edge  of  the  basal  area. 

Deltocephalus  reflexus  Osborn  and  Ball. 

(PI.  XXX,  fig.  6.) 

Length,  1.18  mm.;  greatest  width,  0.09  mm.  About  the  same  width  for  en- 
tire length;  tapers  gradually  caudad  to  tip;  no  preapical  prominence;  slightly 
but  distinctly  curved,  tip  gradually  narrowed,  extreme  apex  sharply  pointed, 
chitinization  light;  strengthening  rod  extends  caudad  as  far  as  tenth  doi"sal 
tooth  from  tip,  nearly  meeting  dorsal  edge.  Toothed  area  on  dorsal  edge  oc- 
cupies somewhat  less  than  the  apical  half  of  the  valve;  teeth  about  forty  in 
number,  rather  small,  of  various  sizes  and  shapes,  presenting  an  irregular, 
crenulate  margin;  a  few  of  the  larger  teeth  bear  secondary  teeth;  the  extreme 
tip  bears  no  teeth ;  ducts  inconspicuous,  invisible  except  for  apices  and  circular 
openings;  open  along  ventral  apical  edge,  along  dorsal  edge  of  toothed  area, 
and  irregularh-  in  the  basal  portion  of  the  valve;  the  two  valves  of  the  pair 
are  joined  one  to  the  other  by  a  poorly  defined,  rather  elongate,  heavily  chit- 
inized connection  present  on  the  dorsal  edge  a  little  past  a  third  of  the  length. 

Deltocephalus  weedi  Van  Duzee. 

(PI.  XXX,  fig.  7.) 

Length,  1.05  mm.;  greatest  width,  0.13  mm.  Narrow  and  rodlike  at  base; 
a  somewhat  wider,  rather  heavily  chitinized  area  extends  almost  to  midpoint ; 
the  apical  half  is  occupied  by  a  still  wider,  less  heavily  chitinized  area  which 
tapers  to  apex;  preapical  prominence  wanting;  slightly  curved,  tip  greatly 
narrowed,  extreme  apex  rounded,  chitinization  moderately  heavy;  heavier  than 
in  D.  reflexus;  strengthening  rod  extends  caudad  as  far  as  or  beyond  last  dor- 
sal tooth.  Toothed  area  on  dorsal  edge  occupies  a  little  more  than  the  apical 
half  of  the  valve;  about  thirteen  large  primary  teeth,  these  are  comparatively 


246  The  University  Science  Bulletin. 

small,  rounded,  fairly  regular  in  size,  shape  and  spacing;  bear  a  few  small 
secondary  teeth  on  caudal  edge ;  margin  of  valve  between  teeth  is  also  notched 
with,  small  secondary  teeth;  tip  notched  with  small  teeth  on  both  dorsal  and 
ventral  edges,  not  continuous  around  the  tip,  nine  present  on  ventral  edge; 
area  of  ducts  conspicuous,  granular  in  appearance,  circular  duct  openings  vis- 
ible; open  along  ventral  apical  edge,  at  extreme  apex,  along  dorsal  edge  of 
toothed  area,  and  irregularly  in  the  basal  area  where  elongate  ducts  are  vis- 
ible; the  two  valves  of  the  pair  are  joined  one  to  the  other  by  an  elongate, 
distinct,  heavily  chitinized  connection  present  on  the  dorsal  edge  about  a  third 
of  the  length. 

Deltocephalus  inimicus  (Say). 

(PI.  XXIII,  fig.  8;  pi.  XXX,  fig.  8.) 

Length,  1.1  mm.;  greatest  width,  0.11  mm.  About  the  same  width  for  en- 
tire length,  narrows  caudad  to  apex,  preapical  prominence  wanting;  slightly 
curved;  tip  greatly  narrowed,  extreme  apex  rounded,  chitinization  moderately 
heavy;  strengthening  rod  extends  caudad  "as  far  as  last  dorsal  primary  tooth. 
Toothed  area  on  dorsal  edge  occupies  apical  half;  primary  teeth  thirteen  in 
number,  small,  rounded,  rather  regular  as  to  size,  shape  and  spacing;  bear  a 
few  small  secondary  teeth  on  caudal  edge;  margin  of  valve  between  primary 
teeth  is  also  notched  with  small  teeth;  ventral  edge  of  tip  is  notched  with 
small  teeth,  dorsal  edge  devoid  of  teeth  at  apex,  fifteen  present  on  ventral 
edge;  area  of  ducts  somewhat  granular,  though  elongate  ducts  are  present, 
circular  duct  openings  visible;  open  along  ventral  apical  edge,  at  extreme 
apex,  along  dorsal  edge  of  toothed  area  and  irregularly  in  the  basal  region ;  the 
two  valves  of  the  pair  are  joined  one  to  the  other  by  an  elongate,  distinct, 
heavily  chitinized  connection  present  on  the  dorsal  edge  at  about  one-third 
the  length. 

Deltocephalus  jlavicosta  Stal. 

(PI.  XXIII,  fig.  9;  pi.  XXX,  fig.  9.) 

Length,  1  mm.;  greatest  width,  0.1  mm.  Apical  portion  bearing  teeth 
occupies  a  little  more  than  one-half  the  length,  is  slightly  wider  than  the 
basal  portion,  narrows  caudad  to  apex;  no  preapical  prominence;  slightly 
curved,  tip  narrowed,  extreme  apex  bluntly  rounded,  chitinization  moderately 
heavy;  strengthening  rod  extends  caudad  as  far  as  last  primary  tooth. 
Toothed  area  on  dorsal  edge  occupies  a  little  more  than  the  apical  half; 
teeth  twelve  to  thirteen  in  number,  small,  rounded,  rather  regular  in  size, 
shape  and  spacing;  bear  a  few  small  secondary  teeth  along  the  caudal  edge; 
margin  of  valve  between  primary  teeth  is  also  notched  with  small  secondary 
teeth;  tip  notched  with  small  teeth  on  ventral  edge,  dorsal  edge  devoid  of 
teeth  at  apex,  eleven  present  on  ventral  edge;  area  of  ducts  somewhat  granu- 
lar in  appearance,  though  elongate  ducts  are  visible;  open  along  ventral  apical 
edge,  at  extreme  apex,  along  dorsal  edge  of  toothed  area,  and  irregularly  in 
the  basal  area;  the  two  valves  of  the  pair  are  joined  one  to  the  other  by  an 
elongate,  curved,  heavily  chitinized  connection  present  on  the  dorsal  edge  at 
about  one-fourth  the  length. 


Readio:    Ovipositors  of  Cicadellid.e.  247 

Deltocephalus  debilis  Uhler. 

(PI.  XXX,  fig.  10.) 

Length,  1.33  mm.;  greatest  width,  0.15  mm.  About  the  same  width  for  en- 
tire length  beyond  curved  basal  connection,  narrows  caudad  to  apex;  pre- 
apical  prominence  wanting;  distinctly  curved,  tip  gradually  narrowed,  extreme 
apex  rovmded,  chitinization  moderatelj^  heavy;  strengthening  rod  extends 
caudad  as  far  as  fourth  dorsal  tooth  from  apex.  Toothed  area  on  dorsal  edge 
occupies  apical  two-fifths  of  length;  primary  teeth  thirteen  to  fourteen  in 
number,  small,  usually  rounded,  irregular  as  to  size,  shape  and  spacing;  for 
the  most  part  bear  no  secondary  teeth,  though  a  few  irregular  secondary 
teeth  are  present ;  the  margin  of  the  valve  between  the  primary  teeth  is 
notched  with  small,  irregular  teeth;  the  tip  is  slightly  and  irregularly  rough- 
ened, but  bears  no  distinct  teeth;  area  of  ducts  conspicuous,  granular  in  ap- 
pearance, though  elongate  ducts  are  visible;  open  along  ventral  apical  edge 
(five),  at  extreme  apex  (one),  along  dorsal  edge  of  toothed  area,  and  in  the 
basal  portion  of  the  valve;  the  two  valves  of  the  pair  are  joined  one  to  the 
other  by  a  distinct,  elongate,  slightly  curved,  heavily  chitinized  connection 
present  on  the  dorsal  edge  of  the  basal  area,  occupying  more  than  one-third 
the  entire  length. 

Deltocephalus  parvulus  Gillette. 

(PI.  XXX,  fig.  11.) 

Length,  0.87  mm.;  greatest  w^idth,  0.08  mm.  About  the  same  width  for 
entire  length  beA^ond  curved  basal  connection,  tapers  gradually  caudad  to 
apex;  preapical  prominence  wanting;  only  slightly  curved,  tip  narrowed,  ends 
in  sharp  point,  chitinization  light ;  strengthening  rod  extends  caudad  almost 
to  last  dorsal  primary  tooth.  Toothed  area  on  dorsal  edge  occupies  a  little  less 
than  the  apical  half;  primary  teeth  nine  to  ten  in  number,  very  small,  rounded, 
fairly  regular  in  size,  shape  and  spacing,  in  most  cases  bear  a  few  secondary 
teeth;  margin  of  valve  between  primary  teeth  also  bears  small,  regular  sec- 
ondary teeth,  which  point  forward;  tip  devoid  of  teeth;  ducts  inconspicuous, 
their  apices  and  circular  openings  alone  being  visible;  open  along  ventral 
apical  edge,  at  extreme  apex,  along  dorsal  edge  of  toothed  area,  and  in  the 
basal  area  where  elongate  ducts  are  visible;  the  two  valves  of  the  pair  are 
joined  one  to  the  other  by  an  elongate,  cui^ved,  distinct,  heavily  chitinized 
connection  present  on  the  dorsal  edge  of  the  basal  area  and  occupying  about 
one-fifth  of  the  entire  length. 

Deltocephalus  collinus  Boheman. 

(PI.  XXX,  fig.   12.) 

Length,  1.3  mm.;  greatest  width,  0.13  mm.  Narrow  and  rodlike  at  base; 
beyond  base  a  broader  and  more  heavily  chitinized  portion  extends  to  mid- 
point; apical  portion  is  still  wider,  less  heavily  chitinized,  tapers  caudad  api- 
cally;  preapical  prominence  wanting;  slightlj'  curved,  tip  greatly  nan-owed, 
extreme  apex  nan-owly  rounded,  rather  blunt,  chitinization  moderately  light; 
strengthening  rod  extends  caudad  as  far  as  seventh  primary  tooth  from  apex. 
Toothed  area  on  dorsal  edge  occupies  a  little  less  than  the  apical  half;  about 
seventeen  primary  teeth  present,  these  are  small,  somewhat  rounded,  very  irreg- 


248  The  University  Science  Bulletin. 

ular  as  to  size,  shape  and  spacing  and  may  or  may  not  bear  a  few  small  second- 
ary teeth  on  the  caudal  edge;  margin  of  valve  between  primary  teeth  notched 
with  small,  irregular  teeth,  especially  caudad  of  apex;  tip  devoid  of  teeth;  area 
of  ducts  conspicuous,  granular  in  appearance,  though  elongate  ducts  are  visible, 
circular  duct  openings  visible;  open  along  ventral  apical  edge  (five),  at  ex- 
treme apex  (one),  along  dorsal  edge  of  toothed  area,  and  irregularly  in  the 
basal  area;  the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a  distinct, 
elongate,  slightly  curved,  heavily  chitinized  connection  present  on  the  dorsal 
edge,  occupying  about  one-fifth  of  entire  length. 

Aconura  argentiolus  (Uhler) . 

(PI.   XXXI,   fig.    9.) 

Length,  1.77  mm.;  greatest  width,  0.11  mm.  About  the  same  width  for  en- 
tire length,  tapers  caudad  to  apex ;  bears  no  i^reapical  prominence ;  slightly 
curved,  tip  narrowed,  rather  blunt,  obtuse-angled  point,  chitinization  very 
light;  strengthening  rod  extends  caudad  to  a  point  between  last  and  next  to 
last  dorsal  tooth.  Toothed  area  on  the  dorsal  edge  occupies  a  little  more  than 
the  apical  half;  teeth  twenty-two  in  number,  very  small,  in  the  general  shape 
of  a  greatly  flattened  obtuse  triangle  with  the  caudal  edge  longer  than  the 
cephalic,  rather  irregular  in  size,  shape  and  spacing;  bear  many  small,  regular 
secondary  teeth  along  both  edges,  more  numerous  on  the  caudal  edge,  continu- 
ous along  entire  dorsal  margin  of  valve  toward  apex;  tip  notched  with  small 
regular  teeth  on  both  dorsal  and  ventral  edges,  not  continuous  around  the  tip, 
teeth  on  ventral  edge  slightly  larger  than  those  or  dorsal ;  areas  of  ducts  incon- 
spicuous; ducts  invisible  except  for  apices  and  circular  openings;  open  along 
ventral  apical  edge,  in  extreme  apex,  along  dorsal  edge  of  toothed  area,  and 
irregularly  in  the  basal  portion  of  the  valve. 

N'ephotettix  cvrtipennis   (Gillette  and  Baker). 

(PI.  XXXI,  fig.   10.) 

Length,  2.5  mm.;  greatest  width,  0.25  mm.  Apical  toothed  half  slightly 
wider  than  basal  portion,  tapers  caudad  to  apex;  bears  an  indistinct,  broadly 
rounded  preapical  prominence  on  ventral  edge;  only  slightly  curved,  tip  nar- 
rowed, ends  in  obtuse-angled  point,  chitinization  moderately  heavy;  strength- 
ening rod  extends  caudad  beyond  last  distinct  primary  tooth.  Toothed  area 
on  dorsal  edge  occupies  the  apical  half;  teeth  twenty-four  to  twenty-five  in 
number,  of  medium  size,  in  the  general  shape  of  a  very  flat,  obtuse  triangle 
with  the  caudal  side  longer  than  the  cephalic,  fairly  regular  in  size,  shape  and 
spacing;  bear  small,  regular,  secondary  teeth  on  both  edges,  continuous  around 
the  apex  of  each  tooth,  seven  to  fifteen  on  caudal  edge  and  one  to  four  on 
cephalic  edge;  tip  notched  with  small  teeth  on  both  dorsal  and  ventral  edges, 
not  continuous  around  the  tip,  about  55  on  ventral  edge,  area  of  ducts  con- 
spicuous, granular  in  appearance  though  elongate  ducts  are  visible,  open  along 
ventral  apical  edge  (five),  at  extreme  apex  (one),  along  dorsal  edge  of  toothed 
area,  and  a  few  in  the  basal  region;  the  two  valves  of  the  pair  are  joined  one 
to  the  other  by  an  elongate,  narrow,  heavily  chitinized  connection  extending 
the  entire  length  on  the  basal  area. 


Readio:    Ovipositors  of  Cicadellid^.  249 

Dnotura  gammaroides  (Van  Duzee). 

(PI.  XXIII,  fig.  10;   pi.  XXXI,  fig.   11.) 

Length,  1.8  mm.;  greatest  width,  0.21  mm.  Apical  half  slightly  wider  than 
basal  half,  tapere  caudad  to  apex;  bears  only  a  suggestion  of  a  preapical  promi- 
nence on  the  ventral  edge;  slightly  curved,  tip  narrowed  with  extreme  apex 
narrowly  rounded,  chitinization  moderately  heavy;  strengthening  rod  extends 
caudad  beyond  last  distinct  dorsal  tooth.  Toothed  area  on  dorsal  edge  occu- 
pies the  apical  half;  teeth  twenty  in  number,  of  medium  size,  in  the  general 
shape  of  a  flat,  obtuse  triangle  with  the  caudal  edge  longer  than  the  cephalic, 
rather  regular  in  size,  shape  and  spacing;  bear  small,  regular  secondary  teeth  on 
both  edges,  continuous  around  the  apex  of  each  tooth,  seven  to  eighteen  on 
caudal  edge,  two  to  se\en  on  ventral  edge;  tip  notched  with  small  teeth  on 
both  dorsal  and  ventral  edges,  practically  continuous  around  the  tip  though 
greatly  reduced  at  extreme  apex;  those  on  ventral  edge  are  larger  and  more 
distinct,  about  fifty-four  present  on  ventral  edge;  area  of  ducts  conspicuous, 
granular  in  appearance,  duct  openings  visible;  open  along  ventral  apical  edge 
(five),  at  extreme  apex  (one),  along  dorsal  edge  of  toothed  area,  and  in-egu- 
larly  in  the  basal  area,  the  two  A'alves  of  the  pair  are  joined  one  to  the  other 
by  an  elongate,  narrow,  heavily  chitinized  connection  present  on  the  dorsal 
edge  of  the  basal  area  for  its  entire  length. 

Tliis  ovipositor  is  very  similar  in  general  appearance  to  that  of  Nephotettix 
curtipennis. 

Genus  Euscelis  BruUe. 

The  ovipositors  of  six  species  of  this  genus  have  been  examined 
and  a  wide  variety  of  forms  found  within  the  genus.  Euscelis  exiti- 
osus  (Uhler)  and  Euscelis  striolus  (Fallen),  both  in  subgenus  Athy- . 
sanus,  are  not  similar  in  any  except  their  grosser  details.  Euscelis 
anthracinus  (Van  Duzee),  in  subgenus  Euscelis,  and  Euscelis  comma 
(Van  Duzee),  in  subgenus  Conomellus,  are  similar  one  to  the  other 
but  not  to  any  other  species  of  the  genus  examined.  Euscelis  cur- 
tesii  (Fitch)  and  Euscelis  bicolor  (Van  Duzee),  both  in  subgenus 
Stirellus,  are  very  similar  one  to  the  other  but  not  to  any  other  spe- 
cies of  the  genus  examined. 

The  ovipositors  of  the  subgenus  Stirellus  are  somewhat  narrower 
in  the  basal  half  than  in  the  apical  half,  which  tapers  to  the  apex 
and  bears  teeth  along  its  dorsal  edge.  The  teeth  differ  in  number 
and  in  the  number  of  secondary  teeth  they  bear.  The  apex  is 
notched  with  small  teeth  present  on  the  ventral  edge  only.  The 
area  of  ducts  is  conspicuous,  granular  in  appearance,  with  the  duct 
openings  visible.  The  two  valves  of  the  pair  are  joined  one  to  the 
other  by  an  elongate,  chitinized  connection  present  on  the  dorsal 
edge  of  the  basal  area. 


250  The  University  Science  Bulletin. 

Euscelis  exitiosus  (Uhler). 

(PI.  XXIII,  fig.  11;  pi.  XXXI,  fig.  1.) 

Length,  2  mm.;  greatest  width,  0.2  mm.  Apical  portion,  which  occupies 
two-thirds  of  length,  is  slightly  wider  than  basal  portion,  tapers  caudad  to 
apex;  bears  a  distinct,  angled,  preapical  prominence  on  ventral  edge;  only 
slightly  curved  beyond  curved  basal  attachment,  rather  abruptly  narrowed  at 
tip,  ends  in  sharp  point,  chitinization  moderately  heavy;  strengthening  rod  ex- 
tends caudad  as  far  as  last  doreal  tooth.  Toothed  area  on  dorsal  edge  occupies 
apical  two-thirds  of  length;  teeth  thirty-four  to  thirty-five  in  number,  rather 
small,  of  a  general  triangular  shape,  with  the  caudal  edge  longer  than  the 
cephalic  and  the  apex  rounded,  fairly  regular  in  size,  shape  and  spacing;  bear 
small  secondary  teeth  on  both  edges,  not  continuous  around  the  apex  of  each 
tooth,  eight  to  eighteen  on  caudal  edge,  two  to  four  on  cephalic  edge;  tip 
notched  with  small,  regular  teeth  on  both  dorsal  and  ventral  edges,  practically 
continuous  around  the  tip,  about  eighty  on  ventral  edge  between  preapical 
prominence  and  extreme  apex;  area  of  ducts  conspicuous,  granular  in  appear- 
ance; open  along  ventral  apical  edge,  along  the  dorsal  edge  of  the  toothed 
area,  one  in  each  tooth,  and  a  very  few  in  the  basal  area;  the  two  valves  of 
the  pair  are  joined  one  to  the  other  by  a  poorly  defined  connection  which  is 
slightly  more  heavily  chitinized  than  the  rest  of  the  basal  area,  upon  whose 
dorsal  edge  it  is  located. 

Euscelis  striolus  (Fallen). 

(PI.  XXIII,  fig.  12;  PL  XXXI,  fig.  2.) 

Length,  LI  mm.;  greatest  width,  0.16  mm.  Apical  portion  occupying  two- 
thirds  of  length  but  little  wider  than  basal  portion,  narrows  caudad  to  tip; 
bears  no  preapical  prominence;  only  slightly  cuiwed,  tip  abruptly  narrowed, 
ends  in  obtuse-angled  point,  chitinization  moderately  heavy;  strengthening 
rod  extends  caudad  as  far  as  second  dorsal  tooth  from  apex.  Toothed  area  on 
dorsal  edge  occupies  apical  two-thirds;  teeth  eleven  to  twelve  in  number, 
rather  large,  of  a  general  triangular  shape  with  the  apices  broadly  rounded, 
fairly  regular  in  size  and  shape,  irregular  in  spacing;  bear  secondary  teeth  of 
various  sizes  and  shapes  on  both  edges,  three  to  nine  on  caudal  edge  and  one 
to  four  on  cephalic  edge;  general  arrangement  of  secondary  teeth  is  very  ir- 
regular; tip  bears  a  few  irregular  teeth  on  the  dorsal  edge  only,  there  is  a 
slight  irregularity  of  the  margin  on  the  ventral  apical  edge;  area  of  ducts  con- 
spicuous, granular  in  appearance,  circular  duct  openings  easily  visible;  open 
along  ventral  apical  edge,  in  extreme  apex,  along  dorsal  edge  of  toothed  area, 
and  in  basal  region  where  many  elongate  ducts  are  visible;  the  two  valves  of 
the  pair  are  joined  one  to  the  other  by  an  elongate,  distinct,  chitinized  con- 
nection present  on  the  dorsal  edge  of  the  basal  area. 

Euscelis  anthracinus  (Van  Duzee). 

(PL   XXXI,   fig.    3.) 

Length,  L4  mm.;  greatest  width,  0.18  mm.  About  the  same  width  for  en- 
tire length  beyond  curved  basal  connection;  preapical  prominence  wanting; 
slightly  curved,  tip  narrowed  by  broadly  rounded  ventral  edge,  extreme  apex 
bluntly  rounded,  cliitinization  rather  heavy;  strengthening  rod  extends  caudad 
as  far  as  last  dorsal  tooth.    Toothed  area  on  dorsal  edge  occupies  apical  half; 


Readio:    Ovipositors  of  Cicadellid^.  251 

teeth  nine  to  ten  in  number,  veiy  small,  rounded,  fairly  regular  as  to  size, 
shape  and  spacing,  being  farther  apart  basally;  bear  a  few  small  secondaiy 
teeth  on  caudal  edge,  margin  of  valve  between  teeth  also  notched  with  small, 
numerous  secondary  teeth;  tip  notched  with  small  teeth  on  both  dorsal  and 
\entral  edges,  not  continuous  around  the  tip,  about  fifteen  present  on  the 
ventral  edge;  area  of  ducts  conspicuous,  granular  in  appearance,  though  elon- 
gate ducts  are  visible;  duct  openings  visible;  open  along  ventral  apical  edge, 
at  extreme  apex,  along  dorsal  edge  of  toothed  area,  and  in  basal  region,  where 
many  elongate  ducts  are  visible;  the  two  valves  of  the  pair  are  joined  one  to 
the  other  by  a  distinct,  elongate,  curved,  heavily  chitinized  connection  present 
on  the  dorsal  edge  of  the  basal  area;  this  connection  bears  a  rounded  promi- 
nence toward  the  caudal  end. 

Euscelis  comma  (Van  Duzee). 

(PI.  XXIII,  fig.  13;  pi.  XXXI,  fig.  4.) 

Length,  1.5  mm.;  greatest  width,  0.22  mm.  About  the  same  width  for  entire 
length  beyond  curved  basal  connection,  tapers  caudad  to  apex;  no  preapical 
prominence;  only  slightly  curved,  tip  narrowed  by  broadly  curving  ventral 
edge,  extreme  apex  narrowly  rounded,  chitinization  rather  heavy;  strengthen- 
ing rod  extends  caudad  to  within  a  short  distance  of  apex.  Toothed  area 
on  dorsal  edge  occupies  a  little  less  than  the  apical  half;  teeth  numerous, 
small,  roimded,  veiy  irregular  as  to  size,  shape  and  spacing  and  present  an 
irregularlj'  crenulate  margin;  tip  bears  no  distinct  teeth  but  is  irregularly 
roughened;  area  of  ducts  conspicuous,  granular  in  appearance,  with  many 
elongate  ducts  visible;  open  along  ventral  apical  edge,  at  extreme  apex,  along 
doi-sal  edge  of  toothed  area,  and  in  the  basal  region  where  many  elongate  ducts 
are  visible;  the  two  valves  of  the  pair  are  joined  one  to  the  other  by  an 
elongate,  distinct,  heavily  chitinized  connection  present  on  the  dorsal  edge  of 
the  basal  area;  it  is  a  little  less  than  half  the  entire  length  and  at  the  point 
of  greatest  width  is  about  one-half  the  width  of  the  valve;  bears  an  indistinct, 
broadly  rounded  prominence  a  little  caudad  of  the  midpoint. 

Euscelis  curtfsii  (Fitch). 

(PI.  XXXI,  fig.  5.) 

Length,  1.1  mm.;  greatest  width,  0.12  mm.  Apical  half  slightly  wider  than 
basal  half,  bears  no  preapical  prominence ;  only  slightly  curved  beyond  curved 
basal  attachment,  gradually  narrowed,  ends  in  narrowly  rounded  extreme  apex, 
chitinization  moderatelj'  light;  strengthening  rod  extends  caudad  as  far  as 
next  to  last  dorsal  tooth.  Toothed  area  on  dorsal  edge  occupies  the  apical 
half;  teeth  fourteen  to  sixteen  in  number,  small,  rounded,  fairly  regular  in  size, 
shape  and  spacing,  though  not  entirely  so;  bear  a  few  secondar.v  teeth  on  the 
caudal  edge  and  an  occasional  secondary  tooth  on  the  cephalic  edge;  tip  notched 
with  small  teeth  on  ventral  edge  onh',  dorsal  edge  devoid  of  teeth  at  extreme 
apex,  ten  present  on  ventral  edge;  area  of  ducts  conspicuous,  granular  in  ap- 
pearance, openings  visible;  open  along  ventral  apical  edge,  at  extreme  apex, 
along  dorsal  edge  of  toothed  area,  and  in  basal  region  where  elongate  ducts  are 
visible;  the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a  distinct, 
elongate,  slightly  curv-ed,  chitinized  connection  present  on  the  dorsal  edge  of 
the  basal  area. 


252  The  University  Science  Bulletin. 

Euscelis  bicolor  (Van  Duzee). 

(PI.  XXXI.  fig.   6;    pi.  XXIII,  fig.   14.) 

Length,  0.92  mm.;  greatest  width,  0.11  mm.  Apical  portion  shghtly  wider 
than  basal  portion,  tapers  caiidad  to  apex;  no  preapical  prominence;  distinctly 
curved,  tip  gradually  narrowed,  ends  in  naiTowly  rounded  extreme  apex, 
chitinization  rather  light;  strengthenuig  rod  extends  caudad  as  far  as  next 
to  last  dorsal  tooth.  Toothed  area  on  dorsal  edge  occupies  apical  two-fifths 
of  length;  teeth  twelve  to  thirteen  in  number,  small,  rounded,  fairly  regular 
in  size,  shape  and  spacing;  bear  a  few  small  secondaiy  teeth  on  caudal  edge, 
margin  of  valve  also  notched  with  small  secondar>'  teeth;  tip  notched  with 
small  teeth  on  ventral  edge  only,  eight  to  ten  in  number;  area  of  ducts  con- 
spicuous, granular  in  appearance,  duct  openings  visible;  open  along  ventral 
apical  edge,  at  extreme  -apex,  along  dorsal  edge  of  toothed  area,  and  in  basal 
region,  where  elongate  ducts  are  visible;  the  two  valves  of  the  pair  are  joined 
one  to  the  other  by  an  elongate  connection  only  slightly  more*  heavily 
chitinized  than  the  rest  of  the  basal  area,  occupying  about  one-sixth  of  the 
entire  length. 

Genus  Eutettix  Van  Duzee. 

The  ovipositors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  generally  similar,  though  possessing  many  differ- 
ences. In  each  case  the  basal  half  is  somewhat  narrower  than  the 
apical  half,  which  narrows  toward  the  apex  and  bears  teeth  along 
its  dorsal  edge.  The  teeth  vary  in  number,  size,  shape,  and  posses- 
sion of  secondary  teeth  in  the  two  species.  The  tip  is  notched  with 
small  teeth  on  one  or  both  edges.  The  area  of  ducts  is  conspicuous, 
granular  in  appearance,  with  the  duct  openings  visible.  The  two 
valves  of  the  pair  are  joined  one  to  the  other  by  a  chitinous  con- 
nection present  on  the  dorsal  edge  of  the  basal  area. 

Eutettix  cinctus  Osborn  and  Ball. 

(PI.  XXIV,  fig.  1;    pi.  XXXI,  fig.  8.) 

Length,  1.5  mm.;  greatest  width,  0.17  mm.  Apical  portion  but  slightly  wider 
than  basal  portion,  narrowed  caudad  toward  apex;  no  preapical  prominence; 
distinctly  curved,  tip  narrowed,  extreme  apex  broadly  rounded,  chitinization 
moderately  heavy;  strengthening  rod  extends  caudad  as  far  as  fourth  dorsal 
tooth  from  apex.  Toothed  area  on  dorsal  edge  occupies  apical  half  of  valve; 
teeth  fourteen  in  number,  of  medium  size,  rounded,  fairly  regular  in  size,  shape 
and  spacing;  bear  secondary  teeth,  one  to  four  on  caudal  edge;  cephalic  edge 
may  also  bear  a  single  secondary  tooth ;  margin  of  valve  is  notched  with  small 
secondary  teeth  between  primary  teeth;  a  few  faintly  visible  teeth  are  present 
on  the  ventral  edge  of  the  tip,  the  dorsal  edge  is  devoid  of  teeth  at  the  extreme 
apex,  about  nine  present  on,;  the  ventral  edge;  area  of  ducts  conspicuous, 
granular  in  appearance,  duct  openings  easily  visible;  open  along  ventral  apical 
edge  (seventeen  to  nineteen),  at  extreme  apex  (three),  along  dorsal  edge 
of  toothed  area,  and  in  the  basal  region  where  elongate  ducts  are  visible;  the 


Readio:    Ovipositors  of  Cicadellid^,  253 

two  valves  of  tho  pair  arc  joined  one  to  the  other  by  a  distinct,  elongate, 
curved,  heavily  chitinized  connection  present  on  the  dorsal  edge  of  the  basal 
area;  bears  a  distinct,  rounded  prominence  about  midway. 

Eutettix  strobi  (Fitch). 

(PI.   XXIV,  fig.   2;    pi.  XXXI,   fig.    7.) 

Length.  1.3  ami.;  greatest  width.  0.17  mm.  Apical  portion  somewhat  wider 
than  basal  portion,  narrows  caudad  to  apex;  preapical  prominence  wanting; 
slightly  curved,  tip  narrowed  by  curving  dorsal  edge,  ends  in  broadly  rounded 
extreme  apex,  chitinization  medium;  strengthening  rod  extends  caudad  as 
far  as  fifth  doi-sal  tooth  from  apex.  Toothed  area  on  dorsal  edge  occupies  a 
little  more  than  the  apical  half;  teeth  twenty-one  to  twenty-four  in  number, 
large,  somewhat  iiregidarly  rounded  with  the  caudal  edge  longer  and  more 
gently  sloping  than  the  cephalic  edge;  bear  small  secondary  teeth  on  both 
edges,  three  to  seven  on  caudal  edge  and  one  to  three  on  cephalic  edge;  tip 
is  notched  with  small  teeth  on  both  dorsal  and  ventral  edges,  not  continuous 
around  the  tip,  fifteen  to  nineteen  on  ventral  edge;  area  of  ducts  conspicuous, 
granular  in  appearance,  duct  openings  easily  visible;  open  along  ventral  apical 
edge  (fourteen),  at  extreme  apex  (one),  along  dorsal  edge  of  toothed  area,  and 
in  basal  region  where  elongate  ducts  are  visible;  the  two  valves  of  the  pair  are 
joined  one  to  the  other  by  a  rather  short,  distinct,  heavily  chitinized  con- 
nection present  on  the  dorsal  edge  in  the  basal  region. 

Genus  Phlepsils  Fieber. 

The  ovipositors  of  three  species  of  this  genus  have  been  examined 
and  found  to  be  generally  similar.  The  ovipositor  in  each  case  is 
about  the  same  width  for  entire  length  and  tapers  caudad  to  the 
apex.  The  toothed  area  on  the  dorsal  edge  occupies  from  a  third 
to  a  half  the  apical  length.  The  primary  teeth  vary  in  number, 
size,  shape  and  number  of  secondary  teeth  in  the  various  species. 
The  tip  is  notched  with  small  teeth  on  both  edges  in  every  case.  The 
area  of  ducts  is  conspicuous,  granular  in  appearance,  with  some 
eloiigate  ducts  visible  and  with  duct  openings  visible ;  the  two  valves 
of  the  pair  are  joined  one  to  the  other  by  an  elongate,  chitinized 
connection  present  on  the  dorsal  edge  of  the  basal  area. 

Phlepsius  spatulatus  Van  Duzee. 

(Pl.  XXIV,  fig.  3;   pi.  XXXII,  fig.  10.) 

Length,  1.9  mm.;  greatest  width,  0.25  mm.  About  the  same  width  for 
entire  length,  tapers  caudad  to  apex;  bears  a  suggestion  of  a  preapical  promi- 
nence on  the  ventral  edge;  slightly  curved,  narrowed  by  curving  ventral  edge, 
rather  bhmtly  rounded  at  extreme  apex,  chitinization  heavy;  strengthening  rod 
extends  cauded  as  far  as  fifth  dorsal  tooth.  Toothed  area  on  dorsal  edge 
occupies  apical  two-fifths;  teeth  fifteen  to  eighteen  in  number,  of  medium 
size,  rounded,  rather  irregular  as  to  size,  shape  and  spacing;  bear  a  few 
irregular  secondary  teeth  on  the  caudal  edge;   cephalic  edge  of  some  teeth 


254  The  University  Science  Bulletin. 

also  bears  a  single  secondary  tooth;  margin  of  valve  between  primary  teeth 
notched  with  small  secondary  teeth;  tip  notched  with  small,  irregular  teeth 
on  both  edges;  area  of  ducts  conspicuous,  granular  in  doi-sal  region,  duct-like 
in  ventral  region,  circular  duct  openings  visible;  open  at  ventral  apical  edge 
(seven),  at  extreme  apex  (one),  along  dorsal  edge  of  toothed  area,  and  in 
basal  region;  the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a 
distinct,  elongate,  heavily  chitinized  connection  present  on  the  dorsal  edge  of 
the  basal  area  and  occupying  about  one-tliird  the  length. 

Phlepsius  excultus  (Uhler). 

(PI.  XXIV,  fig.  i;   pi.  XXXII,  fig.  9.) 

Length,  1.8  mm.;  greatest  width,  0.17  mm.  About  the  same  width  for  entire 
length,  tapers  caudad  to  apex;  bears  a  suggestion  of  a  preapical  prominence 
in  the  ventral  edge,  extreme  apex  rounded,  chitinization  moderately  heavy; 
strengthening  rod  extends  caudad  as  far  as  fourth  dorsal  tooth  from  apex. 
Toothed  area  on  dorsal  edge  occupies  a  little  more  than  the  apical  third  of 
the  length;  teeth  fifteen  to  seventeen  in  number,  rather  small,  roimded, 
fairly  regular  in  size  and  shape  but  uneven  in  spacing;  bear  a  few  secondary 
teeth  on  caudal  edge,  and  a  few  primary  teeth  bear  a  single  secondary  tooth 
on  the  cephalic  edge,  margin  of  valve  between  primary  teeth  bears  small 
secondary  teeth;  tip  notched  with  small  teeth  on  both  dorsal  and  ventral  edges, 
practically  continuous  around  the  tip;  eight  large  teeth  on  ventral  edge;  these 
may  be  simple  or  may  bear  secondary  teeth,  irregular  as-  to  arrangement; 
area  of  ducts  conspicuous,  ducts  elongate  for  the  most  part  though  a  granular 
area  is  present  dorsally,  duct  openings  visible;  open  along  ventral  apical  edge 
(six),  at  extreme  apex  (one),  along  dorsal  edge  of  toothed  area,  and  in  the 
basal  region;  the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a  dis- 
tinct, elongate  heavily  chitinized  connection  present  on  the  dorsal  edge  of  the 
basal  area,  occupying  more  than  a  third  of  the  total  length,  this  connection 
irregularly  roughened  along  its  dorsal  edge. 

Phlepsius  irroratus   (Say). 

fPl.  XXIV,  fig.  5;  pi.  XXXII,  fig.  8.) 

Length,  1.5  mm.;  greatest  wddth,  0.17  mm.  About  the  same  width  for 
entire  length  beyond  curved  basal  attachment,  narrows  caudad  to  apex;  pre- 
apical prominence  wanting;  slightly  curved,  tip  gradually  narrowed,  narrowly 
rounded  at  extreme  apex,  chitinization  moderately  light;  strengthening  rod 
extends  caudad  as  far  as  second  dorsal  tooth  from  apex.  Toothed  area  on 
dorsal  edge  occupies  a  little  less  than  the  apical  half;  teeth  nineteen  to  twenty 
in  number,  of  medium  size,  in  the  general  shape  of  an  obtuse  triangle  with 
caudal  edge  longer  than  the  cephalic  and  the  apex  rounded  broadly,  fairly 
regular  as  to  size  shape  and  spacing,  but  not  entirely  so ;  bear  a  few  secondary 
teeth  on  caudal  edge  and  also  a  few  bear  a  single  secondaiy  tooth  on  the 
cephahc  edge;  tip  is  notched  with  small,  indistinct  teeth  on  both  dorsal  and 
ventral  edge,  not  continuous  around  the  tip,  about  fifteen  on  ventral  edge; 
area  of  ducts  conspicuous,  granular  in  appearance,  duct  openings  easily 
visible;  open  along  ventral  apical  edge  (ten),  at  extreme  apex  (one),  along 
dorsal  edge  of  toothed  area,  and  in  the  basal  region  where  elongate  ducts  are 


Readio:    Ovipositors  of  Cicadellid^.  255 

visible;  the  two  ^■alves  of  the  pair  are  joined  one  to  the  other  by  a  heavily 
chitinized  connection  present  on  the  dorsal  edge  of  the  basal  area,  occupying 
about  a  fourth  of  the  entire  length  and  a  half  of  the  width. 

Genus  Acinopterus  Van  Duzec. 

The  ovipositors  of  three  species  of  this  genus  have  been  examined 
and  found  to  be  very  similar.  In  each  case  the  ovipositor  is  greatly 
curved,  the  basal  portion  only  slightly  narrower  than  the  apical 
portion,  which  bears  teeth  along  its  dorsal  edge.  The  primary  teeth 
are  few  in  number,  and  differ  in  number  and  in  the  number  of  sec- 
ondary teeth  they  bear  in  the  various  species.  The  margin  of  the 
valve  between  the  primary  teeth  is  notched  with  small  teeth.  The 
tip  in  each  case  is  notched  with  small  teeth  on  both  dorsal  and 
ventral  edges.  The  area  of  ducts  may  or  may  not  be  conspicuous, 
when  visible  granular  in  appearance  with  duct  openings  visible. 
The  two  valves  of  the  pair  are  joined  one  to  the  other  by  an  elon- 
gate, curved,  chitinous  connection  present  on  the  dorsal  edge  of 
the  basal  area. 

Acinopterios  acuminatus  Van  Duzee. 

(PI.  XXIV,  fig.  6;   pi.  XXXII,  fig.  3.) 

Length,  1.5  mm.;  greatest  width,  0.12  mm.  Apical  portion  only  slightly 
wider  than  basal  portion,  tapers  slightly  caudad;  bears  a  small  preapical 
prominence  on  the  ventral  edge;  greatly  curved,  chitinization  moderately 
light;  strengthening  rod  extends  caudad  as  far  as  last  or  next  to  last  dorsal 
tooth  of  large  size.  Toothed  area  on  dorsal  edge  occupies  somewhat  less  than 
the  apical  half;  primary  teeth  seven  in  number,  small,  rounded,  regular  as  to 
size  and  shape,  unevenly  .spaced;  bears  three  to  four  small  secondary  teeth 
on  the  caudal  edge,  margin  of  valve  between  primary  teeth  bears  numerous 
small,  regular,  secondary  teeth;  tip  notched  with  small  teeth  on  both  dorsal 
and  ventral  edges,  not  continuous  around  the  tip,  about  fifteen  present  on  the 
ventral  edge;  area  of  ducts  conspicuous,  granular  in  appearance,  circular 
duct  openings  visible;  open  along  ventral  apical  edge  (seven),  at  extreme 
apex  (two),  along  dorsal  edge  of  toothed  area,  and  in  the  basal  region;  the 
two  valves  of  the  pair  are  joined  one  to  the  other  by  a  distinct,  curved,  heavily 
chitinized  connection  present  on  the  dorsal  edge  of  the  basal  area. 

Acinopterits  viridis  Ball. 

(P!.  XXXII,  fig.  1.) 

Length,  1.7  mm.;  greatest  width,  0.16  mm.  Apical  portion  only  slightly 
wider  than  basal  portion,  nan-owed  caudad  at  apex,  bears  no  preapical  promi- 
nence; greatly  curved,  tip  narrowed,  ends  in  rounded  extreme  apex,  chitiniza- 
tion light;  strengthening  rod  extends  caudad  almost  to  apex.  Toothed  area 
on  dorsal  edge  occupies  the  apical  third;  only  two  primary  teeth  present, 
these  located  near  the  base  of  the  toothed  area,  small,  rounded,  bear  two  to 
three  small  secondary  teeth  on  their  caudal  edges;  the  margin  of  the  valve  is 
notched  with  many  small,  regular  secondary  teeth;  tip  is  notched  on  both 


256  The  University  Science  Bulletin, 

edges  with  small  teeth,  continuous  around  the  tip,  about  ten  on  ventral  edge; 
area  of  ducts  inconspicuous,  ducts  invisible  except  for  circular  openings;  open 
along  ventral  apical  edge  (six),  at  extreme  apex  (one),  along  dorsal  edge  of 
toothed  area,  and  in  basal  area;  the  two  valves  of  the  pair  are  joined  one  to 
the  other  by  a  poorly  defined,  elongate  connection  present  on  the  dorsal  edge 
of  the  basal  area,  only  slightly  more  heavily  chitinized  than  the  rest  of  the 
basal  area. 

Acinopterns  angulatus  Lawson. 

(PI.  XXXII,  fig.  2.) 

Length,  1.2  mm.;  greatest  width,  0.12  mm.  Apical  portion  only  slightly 
wider  than  basal  portion,  narrows  slightly  caudad  at  apex;  no  distinct 
preapical  prominence  present;  greatly  curved,  tip  narrowed,  extreme  apex 
rounded,  chitinization  moderately  heavy,  heavier  than  in  A.  acuminatus  and 
A.  viridis ;  strengthening  rod  extends  caudad  as  far  as  next  to  last  primary 
tooth.  Toothed  area  on  dorsal  edge  occupies  a  little  more  than  the  apical 
third;  teeth  eight  in  number,  small,  rounded,  regular  in  size  and  shape,  some- 
what unevenly  spaced;  bear  secondary  teeth  on  caudal  edge,  three  to  foiu-  in 
mmiber;  the  margin  of  valve  between  primary  teeth  is  notched  with  small, 
numerous,  regular,  secondaiy  teeth;  tip  notched  with  small  teeth  on  both 
dorsal  and  ventral  edges,  practically  continuous  around  the  tip,"  about  fifteen 
on  ventral  edge;  area  of  ducts  conspicuous,  granular  in  appearance,  circular 
duct  openings  visible;  open  along  ventral  apical  edge  (five  to  six),  at  ex- 
treme apex  (one),  along  dorsal  edge  of  toothed  area,  and  in  basal  region;  the 
two  valves  of  the  pair  are  joined  one  to  the  other  by  a  distinct,  elongate, 
heavily  chitinized  connection  present  on  the  dorsal  edge  of  the  basal  area. 

Genus  Thamnotettix  Zetterstedt. 

The  ovipositors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  generally  similar.  In  each  case  the  basal  half  is 
distinctly  narrower  than  the  apical  half,  which  narrows  caudad  at 
the  apex  and  bears  teeth  along  its  dorsal  edge.  The  primary  teeth 
differ  in  number,  shape,  and  number  of  secondary  teeth  they  bear 
in  the  two  species.  The  tip  is  notched  with  small  teeth  on  both 
edges.  The  area  of  ducts  is  conspicuous,  granular  in  appearance, 
with  the  duct  openings  visible.  The  two  valves  of  the  pair  are 
joined  one  to  the  other  by  a  heavily  chitinized  connection  present 
on  the  dorsal  edge  of  the  basal  area. 

Thamnotettix  clitellarms   (Say). 

(PI.'  XXXII,   fig.   5.) 

Length,  1.4  mm.;  greatest  width,  0.18  mm.  Apical  portion  plainly  wider 
than  basal  portion,  narrows  caudad  toward  apex;  bears  a  very  small  preapical 
prominence  on  ventral  edge;  slightly  curved,  tip  narrowed,  extreme  apex 
broadly  rounded,  chitinization  medium;  strengthening  rod  extends  caudad  as 
far  as  fourth  dorsal  tooth  from  apex.  Toothed  area  on  dorsal  edge  occupies 
a  little  more  than  the  apical  half;  teeth  twenty  to  twenty-three  in  number,  of 


Re.\dio:    Ovipositors  of  Cicadellid.e.  257 

medium  size,  of  a  general  triangular  shape,  with  the  caudal  edge  longer  than 
the  cephalic  and  the  apex  rounded  broadly,  fairly  regular  as  to  size,  shape, 
and  spacing,  though  not  entirely  so;  bear  small  secondary  teeth  on  caudal 
edge  and  a  few  also  bear  a  single  secondary  tooth  on  the  cephalic  edge;  tip 
notched  with  small  teeth  on  both  edges,  not  continuous  around  the  tip,  those 
on  ventral  edge  larger  and  more  distinct,  about  fourteen  present  on  ventral 
edge ;  area  of  ducts  conspicuous,  granular  in  appearance,  circular  duct  openings 
visible;  present  along  ventral  apical  edge  (twelve),  at  extreme  apex  (one),  along 
dorsal  edge  of  toothed  area,  and  in  basal  region  where  elongate  ducts  are  visible; 
the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a  distinct,  heavily 
chitinized,  rather  short,  rectangular  connection  present  on  the  dorsal  edge  of 
the  basal  area. 

Thamnotettix  longidus  Gillette  and  Baker. 

(PI.  XXXII,  fig.   4.) 

Length,  1.3  mm.;  greatest  width,  0.17  mm.  Apical  portion  distinctlj^  wider 
than  basal  portion,  tapers  caudad  at  apex;  no  preapical  prominence;  slightly 
curved,  tip  narrowed,  extreme  apex  narrowly  rounded,  chitinization  medium; 
strengthening  rod  extends  caudad  as  far  as  sixth  dorsal  tooth  from  apex. 
Toothed  area  on  dorsal  edge  occupies  somewhat  more  than  apical  half;  thirty- 
one  primarj'  teeth  present,  in  the  general  shape  of  an  obtuse  triangle,  of 
medium  size,  fairlj'  regular  as  to  size  and  shape  but  not  entirely  so,  uneven  in 
spacing;  bear  small  secondary  teeth  on  both  edges,  four  to  twelve  on  caudal 
edge  and  one  to  four  on  cephalic  edge,  secondary  teeth  continuous  around  the 
apex  of  each  tooth;  tip  notched  with  small  teeth  on  both  dorsal  and  ventral 
edge,  not  continuous  around  the  apex,  eight  on  ventral  edge;  area  of  ducts 
conspicuous,  granular  in  appearance,  duct  openings  visible;  open  along  ven- 
tral apical  edge  (eight),  at  extreme  apex  (one),  along  dorsal  edge  of  toothed 
area  (about  one  to  each  tooth),  and  in  the  basal  area;  the  two  valves  of  the 
pair  are  joined  one  to  the  other  by  a  distinct,  elongate,  heavily  chitinized 
connection  present  on  the  dorsal  edge  of  the  basal  area,  occupies  less  than  a 
fifth  of  the  entire  length. 

Genus  Chlorotettix  Van  Duzee. 

The  ovipositors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  generally  similar.  The  basal  third  may  or  may  not 
be  narrower  than  the  apical  two-thirds,  which  tapers  to  the  apex 
and  bears  teeth  along  its  dorsal  edge.  The  primary  teeth  differ  in 
number,  shape,  and  number  of  secondary  teeth  in  the  two  species. 
The  tip  is  notched  on  both  edges  with  small  teeth,  not  continuous 
around  the  tip.  The  area  of  ducts  is  granular  in  appearance  and 
the  duct  openings  are  visible.  The  two  valves  of  the  pair  are  joined 
one  to  the  other  by  a  chitinous  connection  present  on  the  dorsal  edge 
of  the  basal  area. 


258  The  University  Science  Bulletin. 

Chlorotettix  spatidatits  Osborn  and  Ball. 

(PI.   XXIV,  fig.   7;    pi.   XXXII,   fig.   6.) 

Length,  1.5  mm.;  greatest  width,  0.24  mm.  Apical  portion  much  wider  than 
basal  portion,  tapers  caudad  at  apex;  preapical  prominence  wanting;  sHghtly 
curved,  tip  rather  abruptly  narrowed,  extreme  apex  narrowly  rounded,  chitin- 
ization  medium;  strengthening  rod  extends  caudad  as  far  as  last  primary 
tooth.  Toothed  area  on  dorsal  edge  occupies  the  apical  two-thirds;  teeth 
twenty-seven  to  twenty-eight  in  number,  of  medium  size,  rather  iiTegular  as 
to  shape,  some  rounded,  some  triangular  and  some  flat  across  the  top;  bear 
two  to  seven  secondary  teeth  on  the  caudal  edge  and  may  or  may  not  bear 
one  to  three  secondary  teeth  on  the  cephalic  edge;  tip  notched  with  small 
teeth  on  both  edges,  not  continuous  around  the  tip,  nine  to  ten  on  ventral 
edge ;  area  of  ducts  conspicuous,  granular  in  appearance  though  many  elongate 
ducts  are  visible,  duct  openings  visible;  open  along  ventral  apical  edge  (eight), 
at  extreme  apex  (one),  along  dorsal  edge  of  toothed  area,  and  in  basal  area; 
the  two  valves  of  the  pair  are  joined  one  to  the  other  by  an  elongate,  rectangu- 
lar, chitinized  connection  present  on  the  dorsal  edge  of  the  basal  area,  only 
slightly  more  heavily  chitinized  than  the  rest  of  the  valve,  occupying  about 
one-sixth  of  entire  length. 

Chlorotettix  galbanatus  Van  Duzee. 

(PI.  XXIV,  fig.  8;   PI.  XXXII,  fig.  7. J 

Length,  1.4  mm.;  greatest  width,  0.21  mm.  About  the  same  width  for  en- 
tire length,  apical  portion  very  slightly  narrower  than  basal  portion,  tapers 
caudad  to  apex;  no  preapical  prominence;  slightly  cun^ed,  tip  gradually  nar- 
rowed, extreme  apex  bluntly  rounded,  chitinization  medium;  strengthening  rod 
extends  caudad  as  far  as  third  dorsal  tooth  from  tip.  Toothed  area  on  dorsal 
edge  occupies  somewhat  less  than  the  apical  two-thirds;  teeth  twenty-eight  in 
number,  of  medium  size,  some  rounded,  others  of  a  general  triangular  shape, 
evenly  spaced;  bear  secondary  teeth,  two  to  eight  on  caudal  edge  and  an  oc- 
casional single  secondary  tooth  on  the  cephalic  edge;  tip  notched  with  small 
teeth  on  both  edges,  not  continuous  around  the  tip,  ten  present  on  ventral 
edge;  area  of  ducts  conspicuous,  granular  in  appearance,  circular  duct  open- 
ings visible;  open  along  ventral  apical  edge  (thirteen),  at  extreme  apex  (one), 
along  dorsal  edge  of  toothed  area,  and  in  the  basal  region,  where  elongate 
ducts  are  visible;  the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a 
distinct,  curved,  heavily  chitinized  connection  present  on  the  dorsal  edge  of 
the  basal  area,  occupying  a  little  less  than  a  fourth  of  the  entire  length. 

Jassus  olitorius  Say. 

(PI.  XXXIII,  fig.  2.) 

Length,  3.7  mm.;  greatest  width,  0.2  mm.  Very  long,  narrow  and  rodlike, 
apical  portion  which  bears  teeth  only  slightly  if  any  wider  than  basal  rod,, 
tapers  only  at  apex;  no  distinct  preapical  prominence  present;  distinctly 
curved,  tip  narrowed,  rounded  at  extreme  apex,  chitinization  moderately 
heavy ;  strengthening  rod  extends  caudad  as  far  as  sixth  dorsal  tooth.  Toothed 
area  on  dorsal  edge  occupies  about  the  apical  fourth;  eleven  teeth  present  on 
each  valve  with  a  large  median  tooth  present  between  teeth  one  and  two; 


Readio:    Ovipositors  of  Cicadellid^.  259 

apical  teeth  veiy  small  and  broadly  rounded,  distal  teeth  larger  and  more 
sharply  pointed;  bear  no  secondary  teeth;  tip  bears  no  teeth;  area  of  ducts 
conspicuous,  ducts  elongate,  rather  straight,  circular  openings  visible;  open 
along  ventral  apical  edge  (twenty-two),  at  extreme  apex  (one),  along  dorsal 
edge  of  toothed  area,  very  noticeably  in  teeth  three  and  four,  in  each  of  which 
three  to  four  ducts  open,  and  in  the  basal  rod;  the  two  valves  of  the  pair  are 
joined  one  to  the  other  by  a  distinct,  elongate,  narrow,  heavily  chitinized  con- 
nection present  on  the  dorsal  edge  of  the  basal  rod. 

Tinobregnius  vittatus  Van  Duzee. 

(PI.  XXXIII,  fig.  1.) 

Length,  2.7  mm.;  greatest  width,  0.19  mm.  Long,  narrow  and  rodlike,  apical 
toothed  portion  only  shghtly  wider  than  basal  rod.  tapers  caudad  at  apex;  no 
preapical  prominence;  distinctly  curved,  tip  evenly  naiTOwed,  extreme  apex 
rounded,  chitinization  moderately  heavy,  heavier  than  in  Jasms  olitorius; 
strengthening  rod  extends  caudad  as  far  as  eighth  dorsal  tooth.  Toothed  area 
on  dorsal  edge  occupies  apical  third;  teeth  ten  to  eleven  in  number,  apical 
teeth  small  and  broadly  rounded,  distal  teeth  larger  and  more  sharply  pointed; 
bear  no  secondary  teeth;  tip  bears  no  teeth;  area  of  ducts  conspicuous,  ducts 
elongate,  rather  straight,  duct  openings  visible;  open  along  ventral  apical 
edges  (twelve),  at  extreme  apex  (one),  along  dorsal  edge  of  toothed  area,  and 
in  the  entire  length  of  basal  rod;  the  two  valves  of  the  pair  are  joined  one  to 
the  other  by  a  distinct,  elongate,  narrow,  heavily  chitinized  connection  present 
on  the  dorsal  edge  of  the  basal  rod,  occupying  about  one-fifth  of  the  length  of 
the  basal  shaft. 

This  ovipositor  is  very  similar  to  that  of  Jassus  olitorius. 

Cicadula  punctifrons  var.  repleta  Fieber. 

(PI.  XXIV,  fig.  9;  pi.  XXXIII,  fig.  3.) 

Length,  2.2  mm.;  greatest  width,  0.17  mm.  About  the  same  width  for  entire 
length,  narrows ■  caudad  at  apex;  no  preapical  prominence;  slightly  curved,  tip 
narrowed  by  curving  ventral  edge,  bears  a  finely  toothed  prominence  on  the 
dorsal  edge,  extreme  apex  narrowly  rounded,  chitinization  very  light;  strength- 
ening rod  extends  caudad  as  far  as  fourth  dorsal  tooth  from  apex.  Toothed 
area  on  doi"sal  edge  occupies  the  apical  two-fifths;  teeth  fifteen  in  number, 
rather  large,  rounded,  regular  as  to  size,  shape  and  spacing;  bear  a  few  fine 
secondary  teeth  on  both  edges,  continuous  around  the  apices  of  the  primary 
teeth,  those  on  caudal  edge  larger  and  more  distinct  than  those  on  ventral 
edge,  two  to  eight  present  on  the  ventral  edge;  there  are  fine,  radiating  hues 
extending  from  the  interior  of  each  tooth  to  the  margin;  the  tip  is  notched 
with  small  teeth  on  both  edges,  not  continuous  around  the  tip,  twenty-five  on 
ventral  edge;  area  of  ducts  inconspicuous,  ducts  invisible  except  for  circular 
openings,  which  are  easily  visible;  open  along  ventral  apical  edge  (five),  at 
extreme  apex  (one),  along  dorsal  edge  of  toothed  area,  and  in  the  basal  area; 
the  two  valves  of  the  pair  are  joined  one  to  the  other  by  a  poorly  defined, 
elongate,  hghtly  chitinized  connection  present  on  the  dorsal  edge  of  the  basal 
area,  occupies  about  one-fourth  the  entire  length. 


260  The  University  Science  Bulletin, 

Genus  Balclutha  Kirkaldy. 

The  ovipositors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  similar.  The  basal  portion  is  somewhat  narrower 
than  the  apical  portion.  The  point  of  greatest  width  is  between 
three-fourths  and  four-fifths  the  length  of  the  valve,  beyond  which 
the  valve  tapers  to  a  very  narrow  apex,  sharply  pointed  or  narrowly 
rounded.  The  valve  bears  only  very  small  teeth  located  at  the  apex. 
The  area  of  ducts  may  or  may  not  be  conspicuous,  is  granular  in  ap- 
pearance with  the  duct  openings  visible.  The  two  valves  of  the  pair 
are  joined  one  to  the  other  by  an  elongate,  lightly  chitinized  connec- 
tion present  on  the  dorsal  edge  of  the  basal  area. 

Balclutha  punctata   (Thunberg). 

(PI.  XXIV,  fig.   10;   pi.  XXXIII,  fig.   4.) 

Length,  0.88  mm.;  greatest  width.  0.1  mm.  Apical  portion  somewhat  wider 
than  basal  portion,  point  of  greatest  width  about  three-fomtlis  the  length,  tapers 
caudad  from  this  point  to  the  apex;  distinctly  curved,  tip  greatly  narrowed, 
extreme  apex  narrowly  rounded,  chitinization  very  light;  strengthening  rod 
extends  caudad  to  within  a  short  distance  of  the  apex.  The  valve  is  toothed 
only  on  its  dorsal  apical  edge  for  about  one-sixth  the  length,  teeth  numerous, 
very  small,  regular;  area  of  ducts  inconspicuous,  faintly  granular  in  appear- 
ance, duct  openings  visible;  open  along  ventral  apical  edge,  at  extreme  apex, 
along  dorsal  apical  edge,  and  in  the  basal  region;  the  two  valves  of  the  pair  are 
joined  one  to  the  other  by  a  lightly  chitinized,  elongate,  narrow  connection 
present  on  the  dorsal  edge  of  the  basal  area. 

Balclutha  impicta   (Van  Duzee). 

(PL  XXIV,  fig.  U;   pi.  XXXIII,  fig.   5.) 

Length,  0.88  mm.;  greatest  width,  0.09  mm.  Apical  portio'n  slightly  wider 
than  basal  portion,  point  of  greatest  width  about  four-fifths  the  length;  only 
slightly  curved,  tip  greatly  narrowed,  extreme  apex  very  sharply  pointed,  chi- 
tinization light,  though  heavier  than  in  B.  punctata;  strengthening  rod  extends 
caudad  almost  to  apex.  The  dorsal  edge  is  toothed  only  for  the  apical  sixth  of 
its  length;  teeth  numerous,  very  small  and  regular;  ventral  edge  also  bears  a 
few  small  teeth  at  the  apex,  farther  apart  than  those  on  the  dorsal  edge,  about 
ten  in  number;  area  of  ducts  conspicuous,  granular  in  appearance,  openings 
visible;  open  along  ventral  apical  edge,  at  extreme  apex,  along  dorsal  apical 
edge,  and  in  the  basal  area;  the  two  valves  of  the  pair  are  joined  one  to  the 
other  by  a  distinct,  elongate,  narrow  connection  present  on  the  dorsal  edge  of 
the  basal  area. 

Eugnathodus  abdominalis  (Van  Duzee). 

(PI.  XXIV,  fig.  12;   pi.  XXXIII,  fig.  6.) 

Length,  0.72  mm.;  greatest  width,  0.09  mm.  Apical  half  slightly  wider  than 
basal  half,  point  of  greatest  width  is  about  three-fourths  the  length,  beyond 
this  point  the  valve  tapers  to  the  apex;  distinctly  curved,  tip  greatly  but 
unevenly  narrowed,  extreme   apex  very   sharply  pointed,   chitinization   very 


Rkadu):    Ovipositors  ok  C'kadkllid.io.  'ilil 

lifjhf ;  .^trrnst honing  rod  extends  caiidad  aliiiosl  to  ajirx.  The  toothed  area  on 
the  dorsal  edge  oecuijie-s  only  the  narrowed  portion  of  the  ai)ex;  these  teeth 
are  very  small,  regular,  nuintM'ous;  the  ventral  edge  bears  no  teeth  at  the  apex, 
but  somewhat  hack  from  the  apex  on  the  widened  portion  the  \entral  edge  is 
notehcd  with  many  fine,  indistinct  teeth;  area  of  ducts  conspicuous,  granular 
in  ajiiiearance.  duct  openings  \  isihie ;  open  along  ventral  apical  edge,  at  ex- 
treme ajicx,  along  dorsal  ajiical  edge,  and  in  basal  region;  there  is  no  evidence 
of  a  chitinous  connection  between  the  two  valves  of  the  i)air. 

This  ovipositor  is  similar  in  appearance  to  the   ovipositors   of  the  genus 
Bnlrlnlha  examined. 

Tribe  Typhlocybini   (  Kirschbaum  I . 
Dikraneura  abnorniis   (Walsh). 

(PI.  XXIV,  fig.  IS:    pi.  XXXIII,  i\g.  7.) 

Length.  0.7  nun.;  greatest  width,  0.08  mm.  Basal  half  narrow  and  rodlike, 
apical  half  wider,  flat,  toothed,  tapers  caudad  to  apex;  basal  portion  greatly 
cm'ved.  ajucal  portion  only  slightly  ciu'ved;  tip  greatly  narrowed,  extreme  apex 
narrowly  rounded,  chitinization  moderately  light;  strengthening  rod  extends 
caudad  as  far  as  sixth  dorsal  tooth  from  apex.  Toothed  area  on  dorsal  edge 
occupies  the  apical  half;  the  two  valves  of  the  pair  are  not  identical  as  to 
teeth,  the  one  having  few  and  the  other  many;  the  one  having  more  bears 
twenty-five  primar.v  teeth,  the.se  rather  small,  in  the  general  shape  of  an  obtuse 
triangle,  fairly  regular  as  to  size  and  shape,  much  reduced  in  size  apically; 
bear  secondary  teeth  along  the  caudal  edges,  three  to  six  in  number;  the  tip  is 
notched  with  small  teeth  on  both  edges,  not  continuous  around  the  tip.  four  on 
the  ventral  edge;  area  of  ducts  inconspicuous,  ducts  visible,  elongate,  rather 
few  in  number;  \isible  openings  present  only  along  dorsal  edge  of  toothed  area 
and  in  basiil  area;  no  distinct  chitinous  connection  between  the  two  valves 
])resent. 

(tonus  Empoasc.\  Walsh. 

The  ovipositors  of  fiv?  species  of  this  genus  have  been  examined 
and  found  to  be  very  similar.  In  this  genus  the  two  valves  of  the 
pair  are  not  identical,  but  differ  in  length,  width,  and  size  and  num- 
ber of  teeth.  The  shorter,  narrower  valve  bears  many  very  small, 
regular  teeth  along  its  dorsal  edge.  The  longer,  broader  valve  bears 
comparatively  few  large  teeth,  which  in  turn  bear  small- secondary 
teeth.  Neither  valve  in  any  of  the  species  examined  bears  teeth  for 
more  than  the  apical  fifth  of  its  length.  Except  in  one  species  the 
tip  of  the  valve  is  notched  with  small  teeth.  The  area  of  ducts  may 
or  may  not  be  conspicuous;  ducts  elongate,  few  in  number. 

Empoasca  trifasciata  Ctillette. 

(PI.  XXXIII,  fig.    HI.) 

Length.  0.8  mm.;  greatest  width.  0.08  nun.  Narrow  and  rodlike,  apical 
toothed  i)ortion  onlj-  slightly  wider  than  basal  shaft;  di.stinctly  curved,  tip  nar- 
rowed, extreme  apex  narrowly  rounded,  chitinization  moderately  light; 
strengthening  rod  extends  caudad  as  far  as  second  dorsal  tooth.     Toothed  area 


262  The  University  Science  Bulletin. 

on  dorsal  edge  occupies  the  apical  fifth  of  the  valve;  the  two  valves  of  the  pair 
differ  as  to  number  and  arrangement  of  teeth ;  the  shorter,  narrower  valve  bears 
many  small,  regular  teeth  along  its  dorsal  edge;  the  longer,  broader  valve  bears 
thirteen  large  primaiy  teeth  along  its  dorsal  edge,  these  rounded,  regular,  and 
bear  one  to  three  secondary  teeth  along  their  caudal  edges;  the  tip  is  notched 
with  small  teeth  on  both  dorsal  and  ventral  edges,  not  continuous  around  the 
tip,  seven  on  \entral  edge  of  valve  bearing  small  teeth,  eleven  on  ventral  edge 
of  valve  bearing  large  teeth;  area  of  ducts  conspicuous,  ducts  easily  visible, 
elongate,  rather  few  in  number;  open  at  apex,  along  dorsal  edge  of  toothed 
area,  and  in  basal  area;  no  distinct  chitinous  connection  present. 

Empoasca  smaragdula  (Fallen). 

(PI.  XXXIII,  fig.  11.) 

Length,  2  mm.;  greatest  width,  0.11  mm.  Narrow  and  rodlike,  about  the 
same  width  for  entire  length,  tapers  caudad  at  apex;  distinctly  curved,  tip 
narrowed,  extreme  apex  narrowly  rounded,  chitinization  medium;  strengthen- 
ing rod  extends  caudad  as  far  as  third  dorsal  tooth.  Toothed  area  on  dorsal 
«dge  occupies  about  the  apical  seventh  of  the  length;  the  two  valves  of  the 
pair  are  not  identical;  the  one  is  shorter,  narrower,  and  bears  only  very  small 
teeth  along  its  dorsal  edge ;  the  longer,  wider  valve  beai-s  thirteen  large  primary 
teeth  along  its  dorsal  edge,  flatly  rounded,  fairly  regular  as  to  size,  shape  and 
spacing;  bear  a  few  indistinct  secondaiy  teeth;  the  tip  of  the  larger  valve  is 
notched  with  small,  indistinct,  irregular  teeth  on  both  dorsal  and  ventral  edges, 
continuous  around  the  apex;  area  of  ducts  conspicuous,  ducts  easily  visible, 
elongate,  few  in  number;  open  at  apex  and  along  basal  shaft;  the  dorsal  edge 
of  the  basal  shaft  is  irregularly  roughened;  no  distinct  chitinovis  connection 
present. 

Empoasca  obtusa  Walsh. 

(PI.  XXIV,  fig.   14:   pi.  XXXIII,  fig.   12  ) 

Length,  0.9  mm.;  greatest  width,  0.05  mm.  Consi.-ts  of  a  narrow,  rodlike 
basal  shaft  and  a  slightly  wider,  toothed  apical  portion  which  tapers  caudad 
to  apex;  distinctly  curved,  tip  only  slightly  narrowed,  extreme  apex  broadly 
rounded,  chitinization  moderately  light;  strengthening  rod  extends  caudad 
as  far  as  second  dorsal  tooth.  The  two  valves  of  the  pair  are  not  identical; 
the  shorter,  narrower  valve  bears  only  very  small,  regular  teeth  along  its 
dorsal  edge;  the  longer,  wider  valve  is  toothed  along  its  apical  seventh  with 
seven  primary  teeth,  these  of  medium  size,  rather  flatly  rounded,  fairly  regular 
as  to  size,  shape  and  spacing,  though  smaller  apically,  and  bear  a  few  irregular, 
indistinct,  secondary  teeth;  no  distinct  teeth  present  on  tip;  area  of  ducts 
rather  conspicuous,  ducts  visible,  few  in  number,  elongate;  open  apically, 
along  dorsaledge  of  toothed  area,  and  along  basal  shaft. 

Empoasca  livingstoni  (iillette. 

(PI.  XXXIII,  fig.  14.) 

Length,  2  mm.;  greatest  width,  0.11  mm.  Narrow  and  rodlike,  about  the 
same  width  for  entire  length,  tapers  caudad  at  apex;  distinctly  curved,  tip 
greatly  narrowed,  extreme  apex  ends  in  acute-angled  point,  chitinization  very 
light;  strengthening  rod  extends  caudad  as  far  as  fifth  dorsal  tooth.  The  two 
valves  of  the  pair  are  not  identical;  the  shorter,  naiTower  valve  bears  only 


Readio:    Ovipositors  of  Cicadellid^.  263 

very  small,  lobular  tooth  for  a  short  distance  on  its  dorsal  edge;  the  longer, 
wider  \alvo  bears  thirteen  primary  teeth  along  the  apical  sixth  of  its  dorsal 
edge,  medium  in  size,  somewhat  triangular  in  shape,  with  the  caudal  edge 
longer  than  tlu^  cephalic,  fairly  regular  in  size,  shape  and  spacing;  bear  small 
secondary  teeth  along  caudal  edge,  three  to  seven  in  number;  tip  of  longer 
valve  notched  with  small  teeth,  continuous  around  the  tip,  thirteen  on  ventral 
edge;  area  of  ducts  inconsiiicuous,  ducts  invisible  except  for  circular  openings; 
open  at  ajiex.  along  dorsal  edge  of  toothed  area,  and  in  basal  shaft. 

Enipoasca  mali  (LeBaron). 

(PI.  XXXIII,  fig.   13.) 

Length.  0.8  mm.;  greatest  width,  0.03  mm.  Very  narrow  and  rodlike,  apical 
toothed  portion  only  very  little  wider  than  basal  shaft;  greatly  curved, 
tip  slightly  narrowed,  extreme  apex  narrowly  rounded,  chitinization  light; 
strengthening  rod  extends  caudad  as  far  as  third  dorsal  tooth.  The  two 
valves  of  the  pair  are  not  identical;  the  shorter,  narrower  valve  bears  only 
very  small,  regular  teeth  along  its  dorsal  edge  for  a  short  distance ;  the  longer, 
wider  valve  bears  eighteen  primary  teeth  along  the  apical  sixth  of  its  dorsal 
edge,  these  of  medium  size,  regular  as  to  size,  shape  and  spacing,  smaller 
apically,  and  bear  a  few  very  fine  and  indistinct  secondary  teeth  on  the 
caudal  edge;  the  tip  of  the  longer  valve  is  notched  with  small  teeth  on  both 
edges,  not  continuous  around  the  tip,  about  seven  on  the  ventral  edge;  area 
of  ducts  inconspicuous,  ducts  invisible  except  for  circular  openings;  open 
at  apex  and  a  few  in  the  basal  shaft. 

Genus  Erythroneura  Fitch. 

The  ovipositors  of  two  species  of  this  genus  have  been  examined 
and  found  to  be  similar  one  to  the  other  and  also  to  the  ovipositors 
of  the  genus  Enipoasca.  In  this  genus  the  two  valves  of  the  pair 
are  not  identical,  but  differ  as  to  length,  width,  and  size  and  number 
of  teeth.  In  one  species  the  small,  more  numerous  teeth  are  borne 
by  the  longer,  wider  valve;  in  the  other  species  the  condition  is 
reversed.  The  teeth  do  not  occur  except  on  the  apical  fourth  of  the 
valve.  The  area  of  ducts  is  inconspicuous,  ducts  when  visible  elon- 
gate, duct  openings  visible. 

Erythroneura  tricincta  Fitch. 

(PI.  XXXIII,  fig.  8.) 

Length,  L2  mm.;  greatest  width,  0.09  mm.  Narrow  and  rodlike,  the  apical 
toothed  portion  only  slightly  wider  than  the  basal  shaft,  tapers  caudad  at 
apex;  distinctly  curved,  tip  narrowed,  extreme  apex  rounded,  chitinization 
very  light ;  strengthening  rod  extends  caudad  to  a  point  about  two-fifths  the 
length  of  the  toothed  portion  of  the  longer  valve.  The  two  valves  of  the 
pair  are  not  identical ;  the  longer,  wider  valve  bears  a  great  many  very  small, 
regular  teeth  along  its  dorsal  edge  for  the  apical  fourth;  the  shorter,  narrower 
valve  bears  slightly  larger,  less  numerous,  regular  teeth  along  its  dorsal  apical 
edge;  tip  notched  with  teeth  on  both  edges,  not  continuous  aroimd  the  tip. 


264  The  University  Science  Bulletin. 

about  twelve  on  the  ventral  edge  of  the  longer  valve;  area  of  ducts  incon- 
spicuous, ducts  invisible  except  for  openings;  open  in  apex  and  along  basal 
shaft. 

-     Erythroneiirn  vulnerata  Fitch. 

(PI.  XXXIII,  fig.  !».) 

Length,  0.8  mm.;  grratett  width,  0.06  mm.  Narrow  and  rodlike,  apical 
toothed  portion  only  slightly  wider  than  basal  shaft,  tapers  caudad  to  apex; 
distinctly  curved,  tij)  narrowed,  extreme  ajiex  rounded,  chitinization  moder- 
ately light;  strengthening  rod  extends  caudad  as  far  as  third  dorsal  tooth. 
The  two  valves  of  the  pair  are  not  identical ;  the  shorter,  narrower  ^-alve  bears 
very  small  teeth  for  a  short  distance  along  its  dorsal  apical  edge;  the  longer, 
wider  valve  bears  ten  jirimary  teeth  along  its  dorsal  edge,  occupying  the  apical 
sixth,  these  of  medium  size,  in  the  general  shape  of  an  obtuse  triangle,  fairly 
regular  in  size,  shape  and  spacing,  bear  two  to  five  small  secondary  teeth  on 
caudal  edge;  tip  notched  with  small  teeth  on  both  edges,  continuous  around 
apex,  about  ten  present  on  ventral  edge  of  longer  valve;  area  of  ducts  in- 
conspicuous, ducts  elongate;   open  apically  and  along  the  basal  shaft. 

CONCLUSIONS. 

An  examination  of  the  descriptions  and  phites  leads  to  several 
conclusions.  The  various  subfamilies  arc  not  distinctly  set  apart 
by  the  characters  of  the  ovipositor.  While  it  is  true  that  in  general 
the  ovipositors  of  the  Bythoscopina^  have  regular,  rounded  teeth; 
those  of  the  Cicadellinse  are  toothed  for  nearly  their  entire  length 
and  have  elongate,  curved  ducts;  those  of  the  Gyponinse  are  stout 
and  heavily  chitinized;  and  those  of  the  Jassinse  have  a  granular 
duct  area  and  a  chitinous  connection  between  the  two  valves  of  the 
pair;  yet  these  characters  are  not  found  in  nil  the  members  of  the 
subfamily  and  are  not  exclusively  found  fii  the  subfamily.  How- 
ever, closely  related  genera  possess  ovipositors  which  are  very  simi- 
lar. Examples  of  this  similarity  ai'e  shown  in  Agalliopsis  and  Ace- 
ratagallia;  Macropsis  and  Oncopsis;  Oncometopia,  Homalodisca  and 
Aulacizes;  Cicadella  and  Graphocephala;  Nephotettix  and  Drio- 
tui'a;  Dorycephalits  and  Hecalus;  Helochara  and  Drcrculacephola; 
Jassus  and  Tinobregrnus;  Balclutha  and  Eugnathodus;  and  Dikra- 
neura,  Empoasca  and  Typhlocyba,  all  in  the  tribe  Typhlocybini, 
which  is  very  clearly  set  apart  by  the  cluiracters  of  the  ovipositor. 

Between  the  species  of  well-defined  genera  there  is  an  indisputa- 
ble generic  similarity.  The  cliaracters  of  size,  chitinization  and 
number  of  teeth  vary,  but  the  characters  of  general  shape,  and  shape 
ahd  arrangement  of  teeth  seem  to  be  constant  within  the  genus. 
Examples  of  generic  similarity  are  shown  in  Idiocerus,  Macropsis, 
Kolla,   Dneculacephala,  Platymetopius,    Deltocephahis,   Balclutha, 


Readio:    ( )\"ir()siTORS  ok  C'icAniiLLiii.K.  265 

Enipoasca,  and  inan>'  ollicrs.  In  tlii'  ;t<linitt('(lly  loose  and  complex 
genus  Eusceli.s  the  ovipositors  show  the  wide  range  of  forms  that 
would  be  cxpeeted,  and  the  same  condition  might  be  found  in  other 
genera  of  equal  complexity. 

Penally,  we  find  constant  characters  of  specific  value  in  the  ovi- 
positor. The  ovipositors  of  seventeen  individuals  of  Cicadella  hi- 
eroglyphica  (Say),  representing  as  wide  a  geographical  range  and 
as  many  color  A'arieties  as  are  to  be  found  in  our  duplicate  collec- 
tion, were  examined  and  found  to  be  constant  in  the  characters 
given,  which  are  sufficient  to  separate  it  from  the  other  species  of  the 
genus  examined.  Several  specimens  each  of  Graphocephala  coc- 
cinea  (Forster)  and  Oncometopia  lateralis  (Fabricius)  were  exam- 
ined in  the  same  way  and  their  characters  also  found  to  be  constant. 
Hence  it  can  safely  be  concluded  that  characters  of  specific  value, 
constant  within  the  range  of  the  species,  are  found  in  the  ovipositors 
of  the  Cicadellidae.  It  is  also  true  that  these  characters  are  accessi- 
ble to  the  general  worker  and  should  not  be  neglected  by  him  in  a 
taxonomic  study  in  this  family. 

BIBLIOGRAPHY. 

1878 — Packard:    Guide  to  the  Study  of  Insects. 

189S-.Marlatt :    Periodical  Cicada.    U.  S.  Dept.  of  Agr.,  Bull.  No.  14.  n.  s. 

1909 — Packard;    Textbook  of  Entomology. 

1910— Stough:    Hackberry  Psylla.  Kansas  U.  Sci.  Bull.,  vol.  V,  No.  9.  p.  121. 

1917 — Van  Duzee:    Catalogue  of  Heniip.  of  N.  A. 

1918 — Xewell:    Comp.  Morph.  of  the  Genitalia  of  In.sects.    Ann.  Ent.  Soc.  Am., 

vol.  XI,  No.  2.  11.  109. 
1919— Kornhauser:    Sexual   Characters  of  Thelia.     Journ.  of  Morph.,  vol.  32. 

Xo.  3.  p.  531. 
1919 — Walker:    Structure  of  Orthoj).  Insects.     Ann.  Ent.  Soc.  Am.,  vol.  XII. 

Xo.  4,  p.  267. 
1920— Lawson:    Cicadellidae  of  Kansas.    Kansas  U.  Sci.  Bull.,  vol.  XII.  Xo.  1. 
1921 — Hilsman:    Ovipositor  of  the  Cicada.    Thesis  MS.,  University  of  Kansas. 


266  The  University  Science  Bulletin. 


PLATE  XXL 

1.  Ventrul   view  of  abdcmen   of  female. 

2.  Dorsal  view  of  abdomen  of  female. 

3.  Cross  section  through  abdomen. 

4.  Bor-^al  view  of  ^^egment  nine,  showing  attachment  of  ovipositor. 

5.  Ventral  view  of  .•segment  nine,  showing  attachment  of  ovipositor. 

6.  Cross  section  through  ovipositor,  showing  relative  position  of  valves. 

7.  Valve  I,  showing  attachment  to  eighth  sternum. 

8.  Detached  iiortion  of  valve  I,  showing  attachment  to  ninth  pleuron. 

9.  Valve   II    (ujijier)    and   valve   I    (lower),  showing   attachment   to  ninth 
sternum. 


Readio:    Ovipositors  of  Cicadellid^. 


267 


PLATE  XXI. 


268  The  University  Science  Bulletin. 


PLATE  XXII. 

1.  Idiucrtns  pallidus  Fitch. 

2.  Agalliupsis  novella   (Say). 

3.  Oncoinetopia  undaia  (Fabricius). 

4.  Oncometopia  lateralis   (Fabricius). 

5.  Homalodisca  triquetra  (Fabricius). 

6.  Aulacizea  irrorata   (Fabricius). 

7.  Cicadella  hieroglyphica   (Say). 

8.  Cicadella  circellata  (Baker). 

9.  Graphocephala  ccccinea  (Forster). 

10.  Helochara  communis  Fitch. 

11.  Drceculacephala  mollipes  (Say). 

12.  Drceculacephala  reticulata  (S'gnoret). 

13.  Kolla  bifida   (Say). 

14.  Kolla  hart  a  (Ball). 


PLATE  XXII. 


10 


(269) 


PLATE  XXIII. 

1.  Gypona  octo-Uncata  (Say). 

2.  Xerophloea  viridis  (Fabricius) . 

■3.  Xestocephalus  pulicariitx  Van  Duzce. 

4.  Scaphoideus  iministus  (Say). 

5.  Platymetopius  acutus  (Say). 

6.  Platymetopius  frontalis  Van  Duzee. 

7.  Mesamia  vitellina  (Fitch). 

8.  Deltoccphalus  inimicus   (Say). 

9.  Deltoccphalus  flavicosta  Stal. 

10.  Driotura  gammaroides  (Van  Duzee). 

11.  Euscelis  exitiosus  (Uhler). 

12.  Euscelis  striolus  (Fallen). 

13.  Eiiscelis  comma  (Van  Duzee). 

14.  Euscelis  bicolor  (Van  Duzee). 

(270) 


PLATE  XXI II. 


««*fes^ 


10 


(271) 


PLATE  XXIV. 

1.  Entettix  cinctus  Osborn  and  Ball. 

2.  Entettix  strobi  (Fitch). 

3.  Phlepsius  spatulatus  Van  Duzee. 

4.  Phlepsius  excultus  (Uhler). 

5.  Phlepsius  irroratus  (Say). 

6.  Acinopterus  acuminatus  Van  Duzee. 

7.  Chlorotettix  spatulatus  Osboin  and  Ball. 

8.  Chlurotettix  galhanatus  Van  Duzee. 

9.  Cicadula  punctijrons  var.  rcpleta  Fieber, 

10.  Balclutha  punctata  (Thunberg). 

11.  Balclutha  impicta   (Van  Duzee). 

12.  Eugnathodus  abdominalis  (Van  Duzee). 

13.  Dikraneura  abnormis  (Walsh). 

14.  Empoasca  obttisa  Walsh. 

(272) 


PLATE  XXIV. 


3 


8 


(273) 


PLATE  XXV. 

1.  AijaUiopsis  novella  (Say). 

2.  Aceratagallia  uhleri  \i\nT>uzee. 

3.  hUocerns  snowi  Gillette  and  Baker. 

4.  Idiocerus  ramentosus  (Uliler). 

5.  Idiocerus  pallidus  Fitch. 

6.  Idiocerus  duzeei  Provanchei'. 

7.  Idiocerus  verticis  (Say). 

8.  Idiocerus  scurra  (Gerniar). 

9.  Idiocerus  nervatus  Van  Duzee. 

10.  BythoHcopus  apicalis  (Osborn  and  Ball). 

11.  Bythoscopus  iniscUus  (Stal). 

(274) 


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27.") 


I'LATE  XXV 


276  The  University  Science  Bulletin. 


PLATE  XXVL 

1.  Macropsis  viiidis  (Fitch). 

2.  Macropis  suturalis  (Osborn  and  Ball). 

3.  Oncopsis  distinctus  (VanDuzee). 

4.  Oncometopia  unda'a  (Fabricius). 

5.  Oncometopia  lateralif^  (Fabricius). 

6.  Homalodisca  triquelra   (Fabricius). 

7.  Aulacizes  irrorata  (Fabricius). 

8.  Cicadella  circellata  (Baker). 

9.  Cicadella  hieroglyphica  (Say). 


Readio:    Ovipositors  of  Cicadellid^. 


277 


PLATE  XXVI. 


278  The  University  Science  Bulletin. 


PLATE  XXVII. 

1.  DrcEculacephala  noveboracensis  (Fitch). 

2.  Drceculacephala  mollipes  (Say). 

3.  Drceculacephala  reticulata  (Signoret). 

4.  Errhomenellus  montaniis  Baker. 

5.  Helochara  communis  Fitch. 

6.  Pagaronia  tripunctata  (Fitch). 


Rkadio:    Ovipositors  of  CicAnKLLin.i': 


279 


PLATE  XXVIT. 


280  The  Ut.'iversity  Science  Bulletin. 


PLATE  XXVIII. 

1.  Kolla  hartii  (Ball). 

2.  Kolla  geometrica  (Signoret). 

3.  Kolla  bifida  (Say). 

4.  Graphocephala  coccinea  (Forster), 

5.  Gypona  bimaculata  Spangberg. 

6.  Gypona  angulata  Spangberg. 

7.  Gypona  pectoralis  Spangberg. 

8.  Gypona  octo-lineata  (Say). 


Readio:    Ovipositors  of  Cicadellid.e. 


281 


PLATE  XXVIII. 


282  The  University  Science  Bulletin. 


PLATE  XXIX. 

1.  Xe^torephahis:  puUcariun  Van  Duzce. 

2.  Xerophloea  viridis  (Fabricius). 

3.  Stroggylovcphalu.s    agrcstis    (Fallen). 

4.  Dorycephalus  platyrhynchns  Osborn. 

5.  Mcmnonia  consobrina  Ball. 

6.  Spangbergiella  mexicana  Baker. 

7.  Parabolocratus  fiavidus  Signoret. 

8.  Hcculus  lineadus  (Uhler). 

9.  Meaamia  straim)ua    (Osborn). 

10.  Mesamia  viiellina  (Fitch). 

11.  Aligia  jiicunda  (Uhler). 


Rkadio:    ()\'ii'081toks  of  ('i('Ai)KLi>in.i':. 


283 


PLATE  XXIX. 


284  The  University  Science  Bulletin. 


PLATE  XXX. 

1.  Scaphoideiis  scalaris  Van  Duzee. 

2.  Scaphoideus  immistus  (Say). 

3.  Platymetopius  acutus  (SajO- 

4.  Plalymetopius  cinereus  Osborn  and  Ball. 

5.  Platymetopius  frontalis  Van  Duzee. 

6.  Deltocephalus  reflcxus  Osborn  and  Ball. 

7.  Deltocephalus  weedi  Van  Duzee. 

8.  Deltocephalus  inimicibs  (Say). 

9.  Deltocephalus  flavicosta  Stal. 

10.  Deltocephalus  debilis  Uhler. 

11.  Deltocephahis  parvulus  Gillette. 

12.  Deltocephalus  collinus  Boheman. 


Readio:    Ovipositors  of  CiCADELLiDyE. 


285 


PLATE  XXX. 


286  The  University  S(;ience  Bulletin. 


PLATE  XXX  r. 

1.  Euscelis  exitiosus  (Uhler). 

2.  Euscelis  striolus   (Fallen). 

3.  Euscelis  anthracinus   (Van  Dnzf c) 

4.  Eusceliji  comma  (Van  Diizee). 

5.  Euscelis  curLesii  (Fitch). 

6.  Euscelis  bicolor  (Van  Duzee). 

7.  Eutettix  strobi  (Fitch). 

8.  Eutettix  cinctus  Osborn  and  Ball. 

9.  Aconura  argenteolus  (Uhler). 

10.  Nephotettix  curtipennm  (Gillette  and  Baker). 

11.  Driotura  garnmaroides   (Van  Duzee). 


Readio:    Ovipositors  of  Cicadellid.e. 


287 


PLATE  XXXI. 


288  The  University  Science  Bulletin. 


PLATE  XXXII. 

1.  Acinopterus  viridis  Ball. 

2.  Acinopterus  angulatus  Lawson. 

3.  Acinopterus  acuminatus  Van  Duzee. 

4.  Thamnotettix  longulus  Gillette  and  Baker. 

5.  Thamnotettix  clitellarius  (Say). 

6.  L'hlorotettix  spatulatus  Osborn  and  Ball. 

7.  Chlorotettix  galbanatv^s  Van  Duzee. 

8.  Phlepsiu^  irroratus  (Say). 

9.  Phlepsdus  excultuji  (Uhler). 

10.   Phlepsius  spatulatus  Van  Duzee. 


Readio:    Ovipositors  of  Cicadellid^. 


289 


PIATE  XXXII. 


290  The  University  Science  Bulletin. 


PLATE  XXXIII. 

1.  Tinohregmus  vittatus  Xnn  Diizee. 

2.  Jassus  olitorius  Say. 

3.  Cicadula  ■punctifrons  var.  repleta  Fieber. 

4.  Balclutha  punctata  (Thunberg). 

5.  Balclutha  impicta  (VanDuzee). 

6.  Eugnathodus  abdominalis  (Van  Diizee). 

7.  Dikrancura  ahnormis  (Walsh). 

8.  Erythroneura  tricincta  Fitch. 

9.  Erythroneura   vulnerata  Fitch. 

10.  Empoasca  trifasciata  Gillette. 

11.  Empoasca  smaragdula  (Fallen). 

12.  Empoasca  obtusa  Walsh. 

13.  Empoasca  mali  (Le Baron). 

14.  Empoasca  livingstoni  Gilli  tie. 


Readio:    Ovipositors  of  Cicadellid.k. 


291 


PLATE  XXXIII. 


INDEX. 

PAOK 

Abdomen  of  the  female,  The 218 

abdominalis,  Eugiuithodus 260 

abnormis,  Dikraneura 261 

Aceratagallia 

uhleri 224 

Acinopterus 255 

acuminatus 255 

angulatus 256 

viridis 255 

Aconura 

argentiolus 248 

Acucephalini 238 

acuminatus,  Acinopterus 255 

acutus,  Platymetopius 243 

Agalliopsis 224 

novella 224 

agrestis,  Stroggj-locephalus 238 

Aligia 

jucunda 240 

angulata,  Gypona 237 

angulatus,  Acinopterus 256 

anthracinus,  Euscelis 250 

apicalis,  Bythoscopus 228 

argentiolus,  Aconura 248 

Aulacizes 

irrorata 230 

Balclutha 260 

impicta 260 

punctata 260 

bicolor,  Euscelis 252 

bifida,  Kolla 232 

bimaculata,  Gypona 237 

Bythoscopinse ; 224 

Bythoscopus 228 

apicalis 228 

miscellus 229 

Chlorotettix 257 

galbanatus 258 

spatulatus 258 

Cicadella 231 

circellata 231 

hieroglyphica 231 

Cicadellinae 229 

Cicadidaj 217 

Cicadula 

punctifrons  var.  repleta 259 

cinctus,  Eutettix 252 

(293) 


294  The  University  Science  Bulletin. 

PAGE 

cinereus,  Platymetopius 244 

circellata,  Cicadella 231 

clitellarius,  Thamnotettix 256 

coccinea,  Graphocephala 234 

collinus,  Deltorephalus 247 

comma,  Euscelis 251 

communis,  Helochara 233 

Conclusions 264 

consobrina,  Memnonia 238 

curtesii,  Euscelis 251 

curtipennis,  Nephotettix 248 

dehilis,  Deltocephalus 247 

Deltocephalus 245 

collinus '. 247 

debilis 247 

flavacosta 246 

inimicus 246 

parvulus 247 

reflexus 245 

weedi 245 

Dikraneura 

abnormis 261 

distinctus,  Oncopsis. 228 

Dorycephalus 

platyrhynchus 239 

Dripculacephala 234 

moUipes 234 

noveboracensis 235 

reticulata 235 

Driotura 

gammaroides 249 

duzeei,  Idiocerus 225 

Empoasca 261 

livingstoni 262 

roali 263 

obtusa 262 

smaragdula 262 

trif asciata 26 1 

Errhomenellus 

montanus 236 

Erythroneura 263 

tricincta 263 

vulnerata 264 

Eugnathodus 

abdominalis 260 

Euscelis 249 

anthracinus 250 

bicolor 252 


Readio:    Ovipositors  of  Cicadellid.^*:.  295 

rA(i  !■; 

comma                                      251 

curtosii 251 

cvitiosiis 250 

strioliis 250 

Eutettix 252 

cinctus 252 

sirol)! ■ 253 

exitiosus,  iMiscelis 250 

cxcultus.  I'hIoi)sius 254 

flavacosta,  Deltooephaius 246 

flavidus,  Parabolocratus 240 

frontalis,  Platymetopius 244 

galbanatus,  Chlorotettix 258 

gammaroides,  Driotura 249 

geometrica,  Kolla 232 

Graphocephala 

coccinea 234 

Uypona 236 

angulata 237 

biraaculata 237 

ooto-lineata 237 

pectoralis 237 

Gyponinae 236 

hartii,  Kolla 233 

Hecalus 

lincatus 239 

Helochara 

communis 233 

hieroglyphica,  Cicadella 231 

Homalodisca 

triquetra 230 

Idiocerus 224 

duzeei 225 

iiervatus 225 

pallidas 225 

ramentosus 226 

.snowi 226 

scurra 226 

verticis 226 

imniistus,  Scaphoideus 243 

impicta.  Balclutha 260 

inimicus,  Deltocephalus 246 

Introduction 217 

irrorata,  Aulacizes 230 

irroratus,  Phlepsius 254 

Jassinae 238 

Jassini 239 


296  The  University  Science  Bulletin. 

Jassus  PAGE 

olitorius 258 

jucunda,  Aligia 240 

Kolla 232 

bifida 232 

geometrica 232 

hartii 233 

lateralis,  Oncometopia 230 

lineatus,  Hecaliis 239 

livingstoni,  Empoasca 262 

longulus,  Thamnotettix 257 

Macropsis 227 

suturalis 227 

viridis 227 

mali,  Empoasca 263 

Memnonia 

consobrina 238 

Mesamia 241 

straminea 241 

vitellina 241 

Methods 223 

mexicana,  Spangbergiella 240 

miscellus,  Bythoscopus 229 

mollipes,  Drseculacephala 234 

montanus,  Errhomenellus 236 

Nephotettix 

curtipennis 248 

nervatus,  Idiocerus 225 

noveboracensis,  Drseculacephala 235 

novella,  Agalliopsis 224 

obtusa,  Empoasca 262 

octo-lineata,  Gypona 237 

olitorius,  Jassus 258 

Oncometopia 229 

lateralis 230 

undata 229 

Oncopsis 

distinctus ' 228 

Oviposition .' 220 

Ovipositor 

Description  of 220 

Taxonomic  use  of 222 

Pagaronia 

tripunctata 236 

pallidus,  Idiocerus 225 

Parabolocratus 

flavidus 240 

parvulus,  Deltocephalus 247 

pectoralis,  Gypona 237 


Readio:    Ovipositors  of  Cicadellid.^:.  297 

PAOK 

Phlepsius 253 

cxcultus 254 

irrorat  us 254 

spatulatus ■.  .  .  253 

Photography  of  ovipositors 218 

Platyinotoi)ius 243 

aculus 243 

cinereus 244 

frontalis 244 

platyrhyiu'hus,  Dorycephalus 239 

pulicarius,  Xestocephalus 239 

punctata,  Balclutha 260 

punctifrons  ver.  repleta,  Cicadula 259 

pygofer 220 

ramentosus,  Idiocerus 226 

reflexus,  Deltocephakjs 245 

repleta,  var.,  Cicadula  punctifrons 259 

reticulata,  Dra'culacephalu 235 

scalaris,  Scaphoideus 242 

Scaphoideus 242 

immistus 243 

scalaris 242 

scurra,  Idiocerus 226 

Segments,  Number  of 218 

smaragdula,  Empoasca 262 

snowi,  Idiocerus 226 

Spangbergiella 

mexicana 240 

spatulatus,  Chlorotettix 258 

spatulatus,  Phlepsius ' 253 

Spiracles,  Number  of 219 

straminea,  Mesamia 241 

striolus,  Euscelis 250 

strobi,  Eutettix 253 

Stroggylocephalus 

agrestis 238 

suturalis,  Macropsis 227 

Thamnotettix ■ 256 

clitellarius 256 

longulus 257 

Tinobregmus 

vittatus 259 

tricincta,  Erythroneura 263 

trifasciata,  Empoasca 261 

triquetra,  Homalodisca 230 

tripunctata,  Pagaronia 236 

Tj'phlocybini 201 

uhleri,  Aceratagallia 224 

undata,  Oncometopia 229 


298  The  University  Science  Bulletin. 

PAGE 

vertiL-is,  Idiocerus 226 

viridis,  Acinopterus 255 

viridis,  Macropsis 227 

viridis,  Xerophloea 238 

vitellina,  Mesamia 241 

vittatus,  Tinobregmus 259 

vulnerata,  Erythroneura 264 

weedi,  Deltocephalus 245 

Xerophloea 

viridis 238 

Xestocephalus 

puliearius 239 


I 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  9— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  9.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

Life  History  Notes  on  Two  Species  of  Saldid.e  (Heteroptera), 

Grace  Olive  Wiley. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE,  KAN. 


Entered  at  tlie  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  9. 


Life  History  Notes  on  Two  Species  of  Saldidse 
(Hemiptera)  Found  in  Kansas.* 

BY  GRACE  OLIVE  WILEY. 

THE  family  characteristics  of  the  saldids  are  so  well  known  that 
a  description  need  not  be  given  here.     A  few  brief  notes  on 
certain  Kansas  species,  however,  may  be  of  interest. 

Nymphs  in  the  third  and  fourth  instars  were  taken  June  1,  1920, 
along  a  little  ravine  leading  to  a  pond  or  pool  which  contained  w^ater 
only  during  the  spring  and  early  summer  when  rains  were  frequent. 
This  little  freshet  and  pond  were  in  a  small  pasture  just  outside  the 
city  limits  of  Chanute,  Kan.  Here,  within  a  few  blocks  of  home,  I 
found  two  different  species  of  saldids,  though  in  not  very  plentiful 
numbers. 

Knowing  that  no  Kansas  saldid  had  yet  been  reared,  I  captured 
all  the  lively  little  fellows  I  could  and  sat  about  the  task  of  rearing 
them.  One  was  a  large  black  species,  the  other  a  much  smaller 
species. 

THE  LARGE  BLACK  SALDID. 
Saldula  major  (Pro\\). 

June  12  there  were  two  adults,  and  by  June  18  the  others  had 
reached  the  adult  state  and  were  mating.  June  21  I  found  twelve 
newly  laid  eggs  on  a  blade  of  grass.  These  hatched  July  3.  With 
what  pleasure  and  interest  these  active  little  fellows  were  watched! 
The  adults  were  shy,  but  very  inquisitive,  and  when  food  was  placed 
in  their  dish  they  were  always  ready  to  run  up  and  take  a  look  at  it, 
curiosity-bent,  it  seemed;  or  when  removing  food  that  had  been 
given  to  them  the  day  previous,  they  were  equally  curious  to  see 
what  was  going  on.  Not  so  with  the  smaller  species;  they  were 
always  trying  to  get  away  or  hide. 

*I  wish  to  thank  Dr.  H.  B.  Hungerford  for  his  help  and  kindly  criticism  and  for  the  loan 
of  a  binocular  from  the  University. 

(301) 


302  The  University  Science  Bulletin. 

The  nymphs  liked  to  stay  hidden  most  of  the  time,  but  would 
come  out  and  feed  quite  readily.  It  was  interesting  to  see  one  of 
the  little  fellows  prodding  around  an  insect  for  a  soft  spot  in  which 
to  insert  its  needlelike  beak.  Nor  does  it  disturb  one  of  them  in  the 
least  to  remove  the  insect  and  place  same  under  the  binocular  to 
watch  the  process  of  feeding. 

The  newly  emerged  nymphs,  either  from  the  egg  or  molt,  are  very 
bright  red  in  color,  with  eyes,  antennae,  beak  and  legs  very  dark. 
They  become  dark,  however,  in  a  very  short  time,  except  the  nymphs 
which  emerge  from  the  eggs;  these  are  red  for  more  than  a  day. 
The  adults  are  very  pale  pink  or  yellowish  when  they  first  emerge, 
with  eyes,  antennae,  beak  and  legs  very  dark.  They  become  dull 
black  in  a  short  time,  with  rather  pale,  obscure  markings  on  the 
wing  covers. 

DESCRIPTION  OF  STAGES. 

See  plate  XXXIV. 
THE  EGG. 

Size.    Length  about  1.2  mm.;  width  across  mdest  part,  .3  mm. 

Color.  Pearly  white  and  transparent  when  first  laid,  becoming  yellow  in 
color  three  days  later.  The  fourth-day  egg  shows  red  eyelike  spot,  and  two 
days  later  the  entire  egg  becomes  red  in  color;  another  day  later,  making  seven 
daj's  from  time  egg  was  laid,  it  is  blood  red  with  verj'  dark,  eyelike  spots. 
When  nine  days  old  the  egg  is  orange  red,  and  on  the  twelfth  day  the  nymph 
emerges.  A  few  hours  before  hatching  the  egg  becomes  pale  or  whitish  in 
color. 

Shape.  General  shape  shown  by  drawing.  It  is  elongate,  larger  and  more 
broadly  rounded  at  one  end,  tapering  and  much  smaller  at  the  other  end,  with 
dorsal  part  arched. 

FIRST  INSTAR. 

Size.  Total  length  of  one  day-old  nymph,  about  1.1  mm.;  width  across 
abdomen,  .6  mm. 

Color.  Red  to  light  reddish  brown,  with  dark  spot  on  fifth  abdominal  seg- 
ment, on  the  median  line  of  the  dorsal  part.  Eyes,  legs,  beak  and  antennae 
black. 

Structural  peculiarities.  Rostrum  reaching  to  the  apex  of  the  posterior  coxae. 
Antennae  four-segmented,  last  joint  darker  in  color  and  larger  than  the  others. 
Tai-si  one-jointed.    Length  of  first  stage,  four  days. 

SECOND   INSTAR. 

Size.    Length  about  1.8  mm.;  width  across  abdomen  about  1  mm. 

Color.  Dark  reddish  brown  on  dorsal  part,^with  ventral  part  red.  Venter 
very  dark  brown,  nearly  black. 

Structural  peculiarities.  Antennae  four-segmented,  with  last  segment  larger 
than  the  other  segments.  Rostrum  reaching  to  the  base  of  the  posterior 
coxae.  Antennae  and  legs  covered  with  very  fine  black  hairs,  some  longer  and 
more  sparsely  placed  than  the  others. 

Shape.    A  trifle  stouter  than  first-instar  nymph. 


Wiley:    Saldid.e.  303 

THUJD    IN  STAR. 

Size.    Lenjrth  from  2.25  to  2.5  inin,;  width  across  abdomen  from  1.3  mm.  to 

1.4  nun. 

Color.  Color  on  the  <lorsum  very  dark  brownish  black,  with  ventral  parts 
brown. 

Shape.    Much  the  same  as  the  second  instar.  only  much  sto\iter. 

FOIKTH    INST.\R. 

Size.    Length.  3.25  mm.  to  3.5  mm.;  width  across  abdomen,  1.9  mm.  to  2  mm. 

Color.  Shining  black  o\er  entire  part ;  most  of  the  ventral  part  black  ex- 
cept the  throat  and  around  the  legs,  which  is  slightly  paler  in  color,  also  the 
anterior  and  lateral  jiarts  of  the  carina  or  keel  of  the  abdomen,  which  is 
whitish  in  color;  venter  very  dark. 

Shape.  Decidedly  stouter  than  third  instar  and  wing  iiads  now  much  in 
evidence. 

Structural  peculiarities.  Rostrum  reaching  to  the  apex  of  the  posterior 
coxae.  Ventral  part  of  the  abdomen  carinated.  Antenna^  long,  rather  stout, 
and  covered,  as  in  the  third  instar.  with  fine  black  hairs. 

FIFTH    IXST.\R. 

Size.  Length  from  about  4.5  mm.  to  4.8  mm.;  width  across  abdomen  at 
widest  part,  2.5  mm.  to  2.625  mm. 

Color.  All  of  dorsum  shining  black  in  color.  Ventral  parts  same  as  in 
fourth  instar,  as  are  also  the  antennae  and  legs.  Second  joint  of  antennae  de- 
cidedly longer  and  more  slender  than  any  of  the  other  joints.  Rostrum  reach- 
ing to  the  base  of  the  posterior  coxiv.  Eyes  emarginate.  Wing  pads  reaching 
to  the  apex  of  the  second  abdominal  segment. 

THE   .ADULT. 

Size.    Female:    length  of  the  body,  not  including  the  antennae  and  legs, 

6.5  mm.  to  6.875  mm.;  width,  2.8  to  3  mm.     Male:    length,  6.25  mm.;  width, 
2.625  to  2  mm. 

Color.  Entire  insect  dull  black  with  a  few  obscure,  pale  markings.  Body 
oblong,  oval.  Ocelli  slightly  apart.  Head  long  and  tapering  to  the  front. 
Eyes  large,  emarginate  and  strongly  exserted.  Scutellum  rather  flat  with  a  de- 
pression in  center  of  disk.  The  posterior  lateral  margins  of  pronotum  and  the 
anterior  margins  of  hemelytra  slightly  refiexed.  Rostrum  long  and  slender, 
first  joint  short  and  thick,  second  joint  very  long,  the  tip  reaching  to  the  base 
of  the  middle  coxae;  third  joint  reaching  to  the  base  of  the  posterior  coxae. 
Color  of  beak  black  and  shining,  with  tip  slightly  paler.  Antennae  very  black, 
stout  and  long;  basal  joint  stout  and  considerably  shorter  than  third  or  fourth 
joints;  second  joint  very  long  and  slender;  last  two  joints  nearly  the  same 
length;  last  joint  slightly  longer  than  the  next  to  last.  Body  covered  with 
sparse,  closeh'  appressed,  golden  pubescence.  Membrane  furnished  with  four 
areas. 

Oviposition.  The  eggs  are  deposited  at  the  base  of  grass  blades, 
or  are  thrust  \vith  the  sharp  ovipositor  through  the  bhndes  of  grass. 
(See  plate  XXXIV.  j 


304 


The  University  Science  Bulletin. 


Incubation.    Twelve  days. 

Maturity.    Adults  come  from  eggs  in  twenty-eight  days. 

General  notes.  Length  of  first  instar,  four  days;  of  second  in- 
star,  three  days;  of  third  instar,  three  days;  of  fourth  instar,  two 
or  three  days;  of  fifth  instar,  four  days.  No  ocelli  are  present  in  any 
of  the  nymphal  stages. 

MEASUREMENTS  IN  MILLIMETERS  OF  NYMPHS  AND  ADULTS  OF  LARGS  SPBCIE=i 

KANSAS  SALDID. 


3 
» 

Body  measurements,  m  m. 

Leg  measurements,  m  ra. 

? 

3 
TO 

>- 
O 

CO 

_3- 

w 

3- 

g 

Length,  antenna .  . 

CO 

$ 

Fore  leg. 

Middle  leg. 

Hind  leg. 

3- 

3 

a; 
so' 

H 
2. 

3 

-1 

St 
5' 

3_ 

Femur 

-• 

1st 

1.1 

.6 

.4 

.5 

.6 

.7 

.3 

.25 

.1 

.35 

.3 

.15 

.4 

4 

.2 

2d 

1.8 

1.0 

.7 

.75 

1.0 

1.1 

.5 

.4 

.2 

.5 

.45 

.25 

.6 

.7 

.3 

3d 

2,5 

1.4 

.9 

1.0 

1.6 

1.6 

.6 

,5 

.25 

.7 

.6 

.3 

.4 

.8 

1.0 

.4 

4th 

3.5 

1.9 

1.1 

1.3 

2.0 

2,0 

.9 

.7 

.3 

1.0 

1.0 

1.2 

1.5 

.6 

5th 

4.8 

2.5 

1.4 

1.9 

2.7 

2.7 

1.2 

1.0 

.5 

1.5 

1.2 

.6 

1.6 

2.2 

.9 

6th  9 

6.5*  6.75t 

2.8 

1.5 

2.0 

3.5 

3.0 

1.5 

1.3 

.7 

1.6 

1.6 

.8 

2.0 

3.0 

1.25 

6th  9 

6.0*  7.0t 

3.0 

1.5 

2.25 

3.5 

3.5 

1.8 

1.4 

.7 

1.8 

1,6 

.7 

2.2 

3.0 

1.25 

6th  c? 

6.0 

2.8 

1.5 

2.0 

3.5 

3.0 

1.5 

1,4 

.7 

18 

16 

.8  1  2.0 

3.0 

1.25 

*  Venter.        t  Elytra. 

These  measurements  were  made  with  micrometer,  Bausch  and  Loml)  binocular.  6.1  oculars,  55  mm.  objec- 
tives. By  placing  millimeter  ruler  under  binocular,  micrometer  measured  ten  lines  to  the  millimeter.  Thus:  10 
lines  micrometer  =  1  mm. 

SMALL  SPECIES  OF  KANSAS  SALDID. 

Saldula  pOlUpes  Fabr.   (?)* 

From  adults  reared  from  nymphs  captured  June  1,  1920,  along 
with  nymphs  of  a  large  black  species  of  saldid,  I  obtained,  June 
23,  several  clear  creamy- white  eggs,  thrust  in  the  stems  and  blades 
of  grass  growing  in  the  jars  in  which  these  saldids  were  confined. 
These  were  not  the  first  eggs  laid,  however,  as  I  also  found  several 
small  nymphs,  but  the  eggs  had  been  so  cleverly  hidden  that  I  had 
not  found  them.  These  eggs  hatched  July  1  to  3,  and  became  adults 
sixteen  days  later.  Eggs  from  this  second  generation  hatched 
August  13  and  August  20,  becoming  adults  also  in  from  sixteen  to 
seventeen  days  from  the  time  the  nymphs  emerged  from  the  eggs. 

On  the  28th  of  July  I  went  to  Texas  to  join  my  husband,  who  was 
in  the  employ  of  an  eastern  oil  company.     I  could  not  think  of 


*Mr.   Hung.-rforcl  says  this  .American  species  is  not   the  same  as  the  Europtan   species. 


Wiley:    8aldid^.  305 

giving  up  tlic  life  histories  i)f  the  two  species  of  saldids  I  was  rearing, 
so  phicing  all  the  little  glasses  containing  them  in  a  covered  basket, 
I  took  them  with  me.  No  doubt  the  people  on  the  train  thought  I 
was  taking  a  lunch  with  me,  or  perhaps  a  pet  dog  or  cat!  I  wonder 
how  many  ladies  would  have  slept  well  that  night  had  they  known 
that  the  basket  contained  live  bugs! 

In  all  the  rearings  I  fed  dead  flies  and  other  soft-bodied  insects,, 
chiefly  mirids  (capsids)  and  cicadellids  (jassids),  as  these  were 
usually  easier  to  obtain  in  large  numbers  either  by  sweeping  or  at 
the  light  at  night. 

DESCRIPTION  OF  STAGES. 

See  Plate  XXXV. 
THE  EGG. 

Size.    Length  about  .6  mm.;  width,  .15  mm. 

Color.  Clear,  creamy-white  when  tirst  laid,  changing  to  yellow.  Duration 
of  egg  stage.  8  to  10  days. 

Shape.  Elongate-cylindrical;  one  end  broadly  rounded  and  considerably 
larger  than  the  other  end.    Dorsal  jiart  arched. 

FIRST   INSTAR. 

Size.  Length  of  body  about  .8  mm.;  width  of  abdomen  at  widest  part, 
.4  mm. 

Color.  Greenish  brown  with  dark  spot  on  the  dorsum  of  abdomen  along 
median  line. 

Structural  peculiarities.  Mostly  head,  with  large,  reddish  eyes;  four-seg- 
mented slender  antennae,  fourth  segment  stouter  than  the  others.  Duration  of 
first  instar,  four  days. 

SECOND  INSTAR. 

Size.    Length,  1.  mm.;  width,  .6  mm. 

Color.  Light  green  with  eyes  and  antennse  dark  reddish  brown.  Dark  spot 
on  the  dorsum  of  abdomen. 

Shape.  Much  the  same  as  first  instar  only  stouter.  Duration  of  instar, 
three  days. 

THIRD   INST.AR. 

Size.    Length,  1.5  mm.;  width,  .8  mm. 

Color.  Head  and  thorax  yellowish;  third  and  fourth  abdominal  segments 
dark  green  with  spot  on  the  dorsum  orange  yellow;  remainder  of  abdomen 
yellowish  green.     Eyes  and  tip  of  antennae  dark  brown. 

Structural  peculiarities.  Tarsi  segmented  as  in  all  the  other  instars;  rostrum 
reaching  to  the  posterior  coxae.    Duration  of  third  instar,  three  days. 

FOLTRTH  INSTAR. 

Size.    Length  of  body,  2  mm.;  width  of  abdomen,  1.2  mm. 

Color.    Grayish  flecked  with  brown. 

Shape.  Nymph  much  stouter  than  in  the  third  instar.  and  wing  pads  reach- 
ing to  the  base  of  the  second  abdominal  segment.  Beak  reaching  to  the  apex 
of  the  posterior  coxae.    Duration  of  fourth  instar,  3  days. 


306  The  University  Science  Bulletin. 

FIFTH  INSTAR. 

Size.  Total  length,  2.7  mm.  to  3  mm.;  width,  1.5  mm.  to  1.6  mm.  in  wides^t 
portion. 

Color.    Grayish  brown  speckled  or  mottled. 

Structural  peculiarities.  Antennae  slender,  fovir-segmented  as  in  the  other 
instars;  fourth  segment  larger  and  stouter  than  the  others;  second  segment 
longest  and  very  slender;  body,  legs  and  antennae  covered  with  spai-se,  short, 
brown  pubescence.  Rostrum  reaching  to  the  apex  of  the  posterior  coxae. 
Wing  pads  reaching  to  base  of  third  abdominal  segment.  Duration  of  fifth 
instar,  three  days. 

THE  ADULTS. 

Size.  Length  of  the  entire  body,  not  including  antennae  and  legs:  Female: 
3.5  mm.  to  tip  of  abdomen;  3,9  to  4  mm.  to  tip  of  hemelytra;  width  of  abdo- 
men across  the  widest  part,  1.6  mm.  to  1.7  mm.  Male:  length  3  mm.  to  tip 
of  hemelytra;  width,  1.5  mm. 

Description.  Oblong-ovate,  black  above,  closely  invested  with  minute  yellow 
pubescence;  eyes  large,  rather  oblong,  brown,  and  very  prominent.  Ocelli 
reddish  and  slightly  apart.  Clypeus  and  tylus  straw  yellow;  rostrum  black 
and  reaching  upon  the  posterior  coxae.  Ventral  part  body  jet  black;  sternum 
and  pectus  closely  appressed  with  silvery-white  pubescence.  Legs  pale  with 
dark  markings.  Pronotum  and  scutellum  rather  flat,  disk  of  pronotum  slightly 
elevated  and  with  a  minute  depression  in  center  of  disk.  Scutellum  a  Uttle 
longer  than  broad.  Hemelytra  brownish  black  with  grayish-white  markings. 
Clavus  with  oblong  spot  at  the  apex.  Corium  with  two  squarish  spots  near 
the  upper,  outer  margin  and  two  very  small  spots  on  the  posterior  margin  near 
membrane.  Embolium  mostly  grayish  white,  there  usually  being  three  dark 
spots  along  the  median  nerve  connected  by  a  dark-colored  line  or  nerve. 
Membrane  gray  with  dark  veins  and  furnished  with  four  areas,  each  having 
one  or  more  smoky  spots  therein.  Antennae  slender,  four-segmented;  basal 
segment  stoutest,  dark  underneath,  pale  above;  second  segment  very  slender 
and  almost  twice  the  length  of  the  first  segment;  third  and  fourth  segments 
about  equal  in  length  and  stouter  than  the  second. 


Wiley:   Saldid.^. 


307 


ME.\SUREMENTS  IN  MILLIMETERS  OF  NYMPHS  AND  ADULTS  OF  SMALL  SPECIES 

KANSAS  SALDID. 


Body  measurements,  mm. 

Leg  measurements,  mm. 

1 

5* 

31 

>• 

1 

a! 

at 

cr 
o 

c 

a- 
a 
o 

Fore  leg. 

Middle  leg. 

Hind  leg. 

D- 

a 

c 

6= 

3. 

3 
c 
-1 

s: 

CO 

i. 

■4 

B 
c 

I 

1st 

.8 

.4 

.350 

.3 

A 

.4 

.125 

.12 

.065 

.15 

.125 

.070 

.16 

.25 

.080 

2d 

1.0  to 
1.1 

.6 

.4 
.5 

.45 
.525 

.5 

.5 
.6 

.2 

,2 

.1 

.3 

.2 

.125 

.3 

.4 

.175 

3d 

1.5 

.8 

.6 

.7 

.8 

.9 

.4 

.35 

.1 

.4 

.35 

.15 

.5 

.6 

.2 

4th 

1.8  to 
2.1 

1. 
1.2 

.6 

.7 

.8 
.9 

1.0 

1.1 

1.2 

.4 
.5 

.4 

.2 

.5 

.6 

.4 
.5 

.2 

.6 

.7 

.7 
.8 

.25 
.3 

5th 

2.7-2.9 
to  3 

1.5 
1.6 

.9 
1.0 

1.25 
1.3 

1.5 

1.5 
1.6 

.7 

.6 

.25 

.8 
.9 

.7 

.3 

.9 
1.0 

1.1 

.5 

6th  9 

3.5*    4.0t 
3.9 

1.6 
1.7 

1.0 

1.3 

1.6 
1.7 

1.6 
1.7 

.9 

.7 

.3 

.9 

.8 

.35 

.4 

1.1 

1.6 

.6 

6thc? 

3.0  to 

elj'tra 

1.5 

.9 

1.2 

1.5 

1.5 

.7 

.6 

.25 

.8 

.7 

.3 

.9 

1.5 

.5 

*  Venter.        t  Elytra. 

Measurements  made  with  micrometer  same  as  in  large  species,    10  lines  micrometer  =  1  millimeter. 


308  The  University  Science  Bulletin. 


PLATE  XXXIV. 
THE  LARGE  SALDID. 

Saldula  major  (Prov.). 

Fig.  1.  Stem  of  grass  with  some  of  the  blades  removed,  showing  egg  of 
saldid. 

Fig.  2.  Under  side  of  grass  blade,  showing  tips  of  eggs  through  the  opening 
made  by  the  sharp  ovipositor. 

Fig.  3.  Upper  side  of  the  same  blade  shown  in  figure  2,  showing  the  eggs 
and  how  they  were  thrust  through  the  leaf. 

Fig.  4.  Egg  in  a  grass  blade  and  part  of  the  blade  torn  away  to  show  the 
tip  of  the  egg  on  under  surface. 

Fig.  5.    The  egg. 

Fig.  6.    Nymph  just  out  of  egg,  still  in  postnatal  molt. 

Fig.  7.    First-instar  nymph. 

Fig.  8.    Second-instar  nymph.  "n 

Fig.  9.    Fourth-instar  nymph. 

Fig.  10.    Third-instar  nymph. 

Fig.  11.    Fifth-instar  njnnph. 

Fig.  12.    Adult  male. 

Fig.  13.    Adult  female,  ventral  view. 

Fig.  14.    Adult  female,  dorsal  view. 


Wiley  :    Saldid.*:. 


309 


PLATE  XXXIV 


310  The  University  Science  Bulletin. 


PLATE  XXXV. 
THE  SMALL  SALDID. 

Saldula  pi/liipes  Fabr.  (?) 

Fig.  1.  The  egg. 

Fig.  2.  First-instar  nymph. 

Fig.  3.  Second-instar  nymph. 

Fig.  4.  Third-instar  nymph. 

Fig.  5.  Fourth-instar  nymph. 

Fig.  6.  Fifth-instar  nymph. 

Fig.  7.  Adult  female. 

Fig.  8.  Adult  male. 


D 


Wiley:    Saldid.e. 
PLATE  XXXV. 


311 


5. 


d 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  10— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  10.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

A  Problem  in  the  Relation  of  Temperature  to  Rate  of  In- 
sect Development  P.  A.  Glenn. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE,  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  10. 


A  Problem  in  the  Relation  of  Temperature  to 
Rate  of  Insect  Development. 

By  p.  A.  GLENN, 

Chief  Inspector,  Division  of  Plant  Industry,  Illinois  State  Department  of 
Agriculture,  Urbana,  111. 

EVEN  the  most  casual  observer  is  familiar  with  the  fact  that  low 
temperatures  inhibit  growth  in  plants  and  animals,  that  hiber- 
nation or  death  will  be  evidenced  by  temperatures  approaching 
freezing,  and  that  high  temperatures  favor  rapid  development.  It 
is  only  in  late  years  that  biologists  have  attempted  to  ascertain 
with  accuracy  the  reaction  of  various  plants  and  animals  to  differ- 
ent degrees  of  heat.  It  has  now  been  quite  definitely  demonstrated 
that  development  will  take  place  only  when  the  temperature  is 
above  a  certain  point,  called  the  zero  of  development,  or  the 
threshold  of  development;  that  within  a  certain  range  of  tempera- 
tures above  this  point  the  increase  in  the  rate  of  development  is 
approximately  proportional  to  the  rise  in  temperature;  and  that 
under  given  conditions  of  high  moisture,  evaporation  and  other 
physical  environmental  factors  aside  from  heat,  there  is  a  tempera- 
ture constant  for  each  period  in  development,  which  is  equal  to  the 
product  of  the  period  by  the  average  temperature  above  the  thres- 
hold of  development. 

This  range  of  temperatures  within  which  the  rate  of  development 
increases  as  the  temperature  rises  is  bounded  at  the  lower  end  by 
the  threshold  of  development,  and  at  the  upper  end  by  what  I 
shall  call  the  degree  of  the  maximum  rate  of  development.  It  is 
found  that  at  temperatures  near  the  lower  limit  of  this  range  the 
rate  of  development  varies  somewhat  faster  than  the  rate  of  change 
in  temperature,  and  at  temperatures  near  the  upper  limit  the  rate 
of  development  varies  somewhat  more  slowly  than  the  rate  of  change 

(317) 


318 


The  University  Science  Bulletin. 


of  temperatures.  These  facts  are  of  importance  to  scientists  who 
wish  to  ascertain  the  exact  relations  between  temperature  and 
development,  but  for  practical  purposes  these  slight  variations 
need  not  be  taken  into  consideration,  especially  when  the  varying 
out-of-door  temperatures  are  to  be  used  as  a  basis  of  study. 

The  problem,  therefore,  dealt  with  in  this  paper  may  be  stated 
as  follows:  Given  the  length  in  days  of  the  period  of  development 
at  different  average  daily  temperatures,  and  the  average  daily 
temperatures  for  each  of  the  periods,  to  find  the  threshold  of  de- 
velopment, the  degree  of  the  maximum  rate  of  development,  the 
temperature  constant,  the  effects  of  temperatures  above  the  de- 
gree of  the  maximum  rate  of  development,  and  how  to  make  cor- 
rections in  the  temperature  factor  when  the  temperature  for  a  part 
of  the  time  during  the  period  was  above  the  degree  of  the  maximum 
rate  of  development. 

The  problem  is  a  simple  one  when  constant  temperatures  are 
used,  but  with  widely  varying  daily  temperatures  as  a  base  for 
study  the  problem  is  more  difficult,  since  the  average  daily  temper- 
atures for  the  periods  must  be  used  and  in  nearly  all  observations 
these  are  influenced  by  temperatures  below  the  threshold  of  develop- 
ment or  temperatures  above  the  degree  of  the  maximum  rate  of 
development. 

In  this  paper  the  method  of  procedure  will  be  illustrated  by  studies 
on  the  pupal  period  of  the  Carpocapsa  pomonella,  based  on  observa- 
tions out  of  doors  on  the  pupal  period  of  3,817  pupae  at  mean  daily 
temperatures  varying  from  52.6°  F.  to  82.6°  F. 

The  following  table  gives  the  complete  results  of  the  study : 


RELATION  OF  TEMPERATURE  TO  LENGTH  OF  PUPAL  PERIOD. 

Temper- 
ature 
range. 

Generation. 

Number 

of 
obser- 
vations. 

Har- 
monic 
average 

of 
period. 

1 
P 

Aver- 
age 
mean 
daily 
temper- 
ature. 

Average  daily 
day-degrees. 

Total  day- 
degrees. 

52-t- 

2(87-F) 

(52-F)- 
2(87-F) 

52+ 

(52+)- 
2(87+) 

52-55 

Hib 

2 
348 
976 
243 
218 
175 
221 
247 
517 
481 
133 
256 

45.5 
35.2 
34.0 

27.7 

13.8 

12.7 

11.5 

10.7 

10.0 

9.4 

9.4 

9.2 

.021 
.028 
.029 
.036 
.072 
.078 
.086 
.093 
.099 
.105 
.106 
.108 

52.6 
55.7 
56.1 
58.5 
69.2 
70.7 
73.1 
74.9 
76.7 
79.1 
80.8 
82.6 

5.1 

6.8 

7.0 

8.6 

17.3 

18.7 

21.1 

22.9 

24.7 

27.1 

28.8 

30.6 

";64' 

.24 

.42 

.88 

1.63 

3.52 

4.12 

5.1 

6.8 

7.0 

8.6 
17.3 
18.7- 
20.8 
22.5 
23.8 
25.5 
25.3 
26.5 

236 
237 
240 
241 
238 
238 
242 
245 
248 
256 
272 
283 

236 

54-55 

Hib 

237 

56-57 

Hib 

240 

58-59 .  .  . 

Hib 

241 

68-69.... 
70-71.... 
72-73.... 
74-75.... 
76-77. . .  . 
78-79 

First  and  second. 
First  and  second. 
First  and  second. 
First  and  second, 
First  and  second, 
First 

238 
238 
240 
241 
239 
240 

80-81 

First 

239 

82-83 

First 

245 

52-83.... 

3,817 

14.4 

.669 

17.3 

.77 

16.6 

251 

240 

Glenn:    Insect  Development.  319 

Tlio  usual  method  for  determining  the  threshold  of  development 
is  to  use  the  reciprocals  of  the  periods  and  the  average  mean  daily 
temperatures  as  the  coordinates  and  plot  the  one  against  the  other, 
then  draw  a  line  which  best  fits  the  points,  and  the  point  where  it 
crosses  the  temperature  axis  is  the  threshold  of  development.  This 
method  serves  quite  well  when  the  temperatures  are  constant,  but 
the  more  widely  the  temperature  varies  during  the  daily  variations 
the  less  accurate  are  the  results.  The  accompanying  figure  is  really  ■ 
four  figures  placed  on  the  same  sheet,  and  represents  the  points 
in  the  various  positions  assumed  by  them  at  the  different  stages 
in  the  solution  of  the  problem. 

The  circles  represent  the  position  of  points  resulting  from  plotting 
the  reciprocals  of  the  periods  against  the  average  mean  daily 
temperatures.  These  points  do  not  fall  in  a  straight  line,  but  at  the 
lower  end  they  are  too  far  to  the  left,  because  the  average  mean 
daily  temperatures  are  the  averages  of  temperature  readings  below 
the  threshold  of  development  (ineffective  temperatures)  as  well  as 
readings  above  this  point,  and  the  points  at  the  upper  end  are  too  far 
to  the  right,  because  the  average  mean  daily  temperatures  here  are 
averages  of  readings  above  the  degree  of  the  maximum  rate  of 
development  (retarding  temperatures)  as  well  as  readings  below 
this  point. 

A  line  that  would  fit  these  points  would  be  too  flat.  Now  the 
points  that  are  least  affected  by  these  ineffective  and  these  retard- 
ing temperatures  are  those  between  the  average  mean  daily  tempera- 
tures of  68  and  72.  A  line  drawn  through  these  two  points  cuts 
the  temperature  axis  at  51.92  degrees.  This  is  so  nearly  52  de- 
gress that,  to  avoid  fractions,  52  was  assumed  to  be  the  approximate 
threshold  of  development. 

The  average  daily  degrees  above  52  were  then  computed  with 
results  as  shown  in  column  7.  The  points  marked  by  dots  were 
plotted  by  using  the  reciprocals  of  the  periods  and  the  average 
daily  temperatures  above  52  degrees.  It  will  be  seen  that  at  the 
lower  end  of  the  line  the  points  come  into  line  with  the  two  points 
used  to  establish  the  line,  but  at  the  upper  end  of  the  figure  they 
are  in  the  same  position  as  the  points  located  by  plotting  reciprocals 
against  average  mean  daily  temperatures. 

It  was  necessary  now  to  ascertain  the  degree  of  the  maximum  rate 
of  development.    The  only  way  to  determine  this  point  is  by  trial. 

It  was  assumed  at  first  that  temperatures  above  the  degree  of  the 
maximum  rate  of  development  were  ineffective  and  did  not  ac- 


320 


The  University  Science  Bulletin. 


'^ 


--4         •' 


^4      ,^ 


Glenn:    Insect  Development.  321 

celerate  the  rate  of  development.  On  this  supposition  84,  85,  86, 
87,  88  and  89  were  taken  in  turn  as  the  degree  of  the  maximum  rate 
of  development  and  the  proper  corrections  made.  This  did  not  bring 
the  points  back  into  line,  as  the  points  higher  up  still  stood  too  far 
to  the  right.  It  was  then  assumed,  after  a  careful  inspection  of  the 
data,  that  temperatures  above  the  degree  of  the  maximum  rate  of 
development  not  only  did  not  accelerate  but  retarded  the  rate  of 
development  in  the  same  proportion  as  an  equal  fall  in  temperatures 
below  this  point  would  retard  it. 

On  this  supposition  84,  85,  86,  87,  88  and  89  degrees  were  in  turn 
taken  as  the  degree  of  the  maximum  rate  of  development,  and  cor- 
rections made  in  the  location  of  the  points  in  each  case.  It  was 
found  that  87  degrees  gave  the  best  results.  Points  represented  by 
the  crosses  were  then  plotted,  using  the  reciprocals  of  the  periods 
and  the  average  daily  temperatures  above  52  degrees,  diminished  by 
twice  the  average  daily  temperatures  above  87  degrees.  This 
brought  all  the  points  nearly  in  line  with  the  two  original  points 
used  in  determining  the  line,  indicating  that  the  second  supposition 
relative  to  the  effects  of  temperatures  above  the  degree  of  the 
maximum  rate  of  development  is  correct  and  that  87  degrees  is  ap- 
proximately the  degree  of  the  maximum  rate  of  development. 

The  corrections  for  temperatures  above  87  degrees  are  entered  in 
column  8  and  the  corrected  average  effective  temperatures  (day- 
degrees)  are  entered  in  column  9. 

The  day-degree  is  used  as  the  thermal  unit  and  is  equivalent  to 
a  temperature  of  one  degree  lasting  for  one  day.  The  product  of 
the  day-degrees  above  52  degrees  (column  7)  by  the  periods  (column 
4)  are  entered  in  column  10.  It  will  be  noted  that  they  are  nearly 
constant  for  the  lower  temperature,  but  increase  as  the  higher  tem- 
peratures are  reached. 

The  product  of  the  day-degrees  above  52  degrees  diminished  by 
twice  the  day-degrees  above  87  degrees  (column  9)  are  recorded  in 
column  11.  It  will  be  seen  that  they  are  nearly  constant  for  all 
temperatures. 

The  average  of  these  products  is  240. 

The  formula  is  C  =  P  (T  —  2t')  in  which  C  =  constant.  P  =  pe- 
riod in  daj's,  T  =  average  day-degrees  above  52  degrees  and  t  = 
average  day-degrees  above  87  degrees.  This  formula  is  the  equation 
of  an  hyperbola.  Plotting  the  periods  expressed  in  days  against  the 
effective  day-degrees  (T  —  2t'),  we  have  the  points  as  represented 


322  The  University  Science  Bulletin. 

by  the  concentric  circles.  The  curve  through  these  points  is  an 
hyperbola  whose  equation  is  P(T  —  2t')  =  C  =  240. 

This  constant  (240)  is  the  average  of  3,817  observations.  We 
may,  therefore,  conclude  that  it  requires  an  average  accumulation 
of  240  effective  day-degrees  to  bring  the  codling  moth  through  the 
pupa  stage.  The  observed  variations  from  this  average  in  the  case 
of  individuals  were  very  great,  due  in  some  cases,  no  doubt,  to 
errors  in  observation  and  in  part  to  individual  differences,  but  for 
the  most  part  to  the  fact  that  the  day  was  used  as  the  unit  of 
time.  By  using  the  day  as  the  unit  of  time  the  actual  period  might 
in  some  cases  be  nearly  a  day  shorter  than  the  observed  period,  and 
in  other  cases  nearly  a  day  longer. 

In  midsummer  the  daily  accumulation  of  effective  day-degrees 
was  sometimes  as  high  as  27,  so  that  the  accumulation  of  day- 
degrees  for  some  of  the  observed  periods  might  be  27  day-degrees 
greater  or  less  than  for  the  actual  period,  or  the  average  240. 

The  following  table  shows  the  range  of  variations: 

Number  of  individuals.  Day-degrees. 

1 166 

1 : 185-194 

4 195-204 

3 205-214 

110 215-224 

602 225-234 

2238 235-244 

653 244-255 

170 254-265 

18 264-275 

7 274-285 

7 284-295 

2 294-305 

Recorded  accumulations  of  less  than  205  or  more  than  275  were 
probably  due  to  errors  in  observations;  if  we  add  27  to  the  former 
and  subtract  27  from  the  latter,  we  still  have  a  variation  of  from 
232  to  248  due  to  individual  differences,  humidity  or  other  causes. 
The  equation  for  the  pupal  period  may,  therefore,  be  written  as 
follows : 

p  (T  — 2t')=240±  8. 

This  paper  was  prepared  merely  to  illustrate  the  method  followed 
in  determining  the  time-temperature  factors,  or  the  equation  for 
the  period.  The  method  is  applicable  to  the  investigation  of  any 
stage  of  an  insect. 

If  the  threshold  of  development  and  the  degree  of  the  maximum 
rate  of  development  should  be  found  to  be  the  same  for  each  of 


Gli:.\.\;    Ln sect  Development.  323 

the  stages  of  a  given  inseet,  the  constant  of  the  equation  for  the 
wliok^  period  fi-oni  the  deposition  of  the  egg  to  tlie  emergence  of 
the  adult  may  be  found  by  adding  the  constants  of  the  equations 
of  the  stages,  and,  by  making  suitable  allowance  for  the  time  which 
elapses  between  the  emergence  of  the  adult  and  the  deposition  of 
eggs,  a  formula  for  the  whole  life  cycle  of  the  insect  may  be  de- 
termined. 

These  equations  may  be  of  i)ractical  value  in  several  ways. 

By  computing  the  normal  daily  effective  day-degrees  in  any  lo- 
cality, the  number  of  generations  of  the  insect  in  that  locality,  and 
the  normal  dates  on  which  the  first  eggs,  or  larvae,  may  be  expected 
to  appear  in  that  place  can  be  ascertained.  If  any  part  of  the 
season  should  be  abnormally  cold  or  warm,  the  amount  of  retard- 
ation or  acceleration  in  development  can  be  computed  by  keeping 
a  daily  record  of  the  effective  day-degrees  and  comparing  them 
with  the  normal  temperatures.  In  this  way  we  should  be  able  to 
ascertain  when  the  injurious  phase  of  any  insect  pest  is  approach- 
ing long  enough  beforehand  to  enable  us  to  take  whatever  pre- 
cautionary measures  are  necessary  to  avoid  injury. 

In  the  case  of  the  codling  moth,  the  dates  when  the  larvae  of  each 
generation  may  be  expected  to  enter  the  fruit  can  be  determined 
long  enough  beforehand  to  enable  the  owners  to  apply  the  spray 
at  the  right  time. 

How  well  this  method  of  forecasting  insect  injuries  will  work  out 
in  practice  only  time  can  tell ;  however,  the  plan  seems  to  be  a 
feasible  one.  and  one  worth  investigating. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  11— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  11.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

Some  Biological  Notes  on  Philippine  Zoology, 

F.  X.  Williams. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4622 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  11. 


Some  Biological  Notes  on  Philippine  Zoology. 

BY  FRANCIS  X.  WILLIAMS. 

THESE  few  notes  are  very  fragmentary  and  not  all  the  data  are 
derived  from  original  investigations.  They  are  set  forth,  how- 
ever, in  hopes  that  the  reader  who  has  not  visited  the  tropics  will 
find  something  of  interest  in  them.  The  writer  has  spent  the  total 
of  about  two  years  in  the  Philippines,  chiefly  at  Los  Baiios,  Luzon, 
and  thus  has*  become  acquainted  with  some  features  of  the  natural 
history  of  this  very  rich  locality. 

ANTS. 

It  is  quite  evident  that  the  ant  is  well  off  in  the  tropics.  An 
abundance  and  variety  of  food,  as  well  as  great  diversity  of  condi- 
tions, suit  the  needs  of  the  multitude  of  its  species  and  make  this 
family  of  insects  a  most  conspicuous  one.  Many  ants  are  harmless, 
or  even  beneficial,  while  others  are  a  great  pest  in  the  house  or  in 
the  field. 

The  well-known,  red  tree  ant,  (Ecophylla  smaragdina  Fab.  (Cam- 
ponotinse)  is  occasionally  a  decided  nuisance  along  the  outskirts  of 
the  forest  and  in  portions  of  cultivated  districts  near  by.  This  ag- 
gressive insect  has  at  least  one  thing  in  its  favor,  for  while  it  bites 
viciously,  it  cannot  sting.  It  is  a  tolerably  large,  long-legged  crea- 
ture that  lives  on  trees  and  bushes,  where  it  constructs  an  ample 
ball-like  nest  of  leaves  spun  together  with  the  silk  of  its  larvae, 
which  are  carried  about  by  the  workers  as  this  silk  is  needed. 
(Ecophylla  frequently  swarms  on  tree  trunks  and  foliage,  and  so  a 
passer-by  may  unknowingly  gather  some  up;  of  these  a  few  will 
eventually  and  unnoticed  gain  a  point  of  vantage  on  his  person,  as 
the  back  of  the  neck,  and  there  bestow  a  vicious  and  startling  nip. 
Certain  homopterous  bugs  that  produce  honeydew  are  attractive  to. 

(329) 


330  The  University  Science  Bulletin. 

these  ants  and  protected  by  them,  but  other  insects  that  are  not 
thus  useful,  and  that  can  put  up  a  struggle,  are  subjected  to  a  gruel- 
ling treatment  before  they  succumb.  And  so  it  is  not  unusual  to 
come  upon  some  unfortunate  bee  or  beetle  stretched  out,  more  or 
less  tautly,  as  fire  fighters  hold  a  life  net,  by  a  circle  of  these  ants, 
each  steadily  and  relentlessly  pulling  on  a  leg  or  other  portion  of 
the  victim's  anatomy. 

The  large  genus  Polyrhachis  is  best  represented  in  the  forest. 
These  ants  also  use  silk  in  nest  building,  but  usually  mix  in  with  it 
small  debris  of  various  sorts.  A  large  number  of  the  species  have  a 
glossy  appearance.  Some  make  a  sort  of  ball-like  nest  among  the 
leaves  of  bushes ;  others  will  cover  over  a  hollow,  as  the  cut  end  of 
a  bamboo,  with  a  sheet  of  debris.  Others  still,  often  large  species, 
will  build  on  the  trunks  of  trees,  where  the  nests  are  placed  in 
crevices  or  among  the  exposed  roots.  In  these,  the  tubular  aperture 
to  the  nest,  with  its  soft,  flexible  rim,  was  found  to  bear  fine,  more 
or  less  inward-projecting  hairs.  Other  nests  may  be  high  up  upon 
tree  trunks,  and  measure  as  much  as  a  foot  and  a  half  long  by  about 
half  as  wide.  They  consist  of  a  bulging  sheet  or  curtain  of  silk,  etc., 
secured  along  its  margin  to  the  tree  trunk.  I  remember  such  a  nest 
on  the  underhanging  side  of  a  tree  with  pale  bark,  and  which  color 
it  somewhat  resembled.  As  the  name  implies,  these  insects  are 
armed,  more  or  less,  and  sometimes  in  quite  a  fantastic  manner,  with 
spines  or  hooks,  which,  as  they  will  often  stick  into  one's  fingers, 
may  make  capture  rather  awkward.  Polyrhachis  is  only  a  fair 
nipper,  but  the  offensives  and  defensives  are  effected  mainly  by 
raising  itself  upon  its  legs,  curving  the  abdomen  under  and  forward, 
and  the  squirting  out  a  fluid. 

The  myrmicine  ants  of  the  genera  Solenopsis  and  Phidologeton 
can  be  very  annoying  insects,  particularly  during  the  wet  season. 
Solenopsis  geminata  Fab.,  also  known  as  the  fire  ant  because  of  its 
smarting  sting,  is  widely  distributed  in  the  tropics  of  both  hemi- 
spheres. It  is  a  rather  small,  yellowish-brown  species  that  often 
swarm  in  the  lowlands.  In  Phidologeton  we  have  a  genus  of  few 
species  that  appear  more  shade-loving  than  Solenopsis  and  which 
also  live  in  very  large  colonies.  The  great  majority  of  specimens  in 
a  nest  are  of  small  size,  but  the  queens  and  the  largest  neuters  or 
soldiers  are  comparatively  immense.  The  ants  are  great  travelers, 
and  their  irregular  columns,  often  encountered  crossing  a  path,  are 
sparsely  though  conspicuously  sprinkled  with  the  great  polished 
soldiers.    These  big,  lumbering  creatures  are  not  to  be  feared;  it  is 


Williams:   Philippine  Zoology.  331 

the  army  of  little  workers  that  must  be  respected,  for  they  are  of  a 
vindictive  nature  and  lose  no  opportunity  of  using  their  sharp 
mandibles.  Phidologeton  will  occasionally  invade  houses  here,  and 
once  in  a  while  some  of  the  occupants  are  driven  out  of  bed.  There 
is  a  record  of  a  village  in  India  having  to  shift  because  of  these 
troublesome  ants  ("Fauna,  British  India,  Hymenoptera,"  II,  p.  161, 
1903;  Bingham  quotes  Rothney).  The  genus  belongs  to  the  Indo- 
Malayan  region,  and,  like  Sole?idpsis  geminata,  is  largely  granivor- 
ous.  It  is  to  be  noted,  then,  that  meat  eaters  are  not  always  the 
fiercest.  Pogonomyrmex,  the  aggressive  and  efficient  stinger  among 
ants  in  some  of  the  more  arid  portions  of  the  United  States,  is  also  a 
granivorous  insect. 

While  the  ponerine  ants  are  comparatively  few  and  inconspicuous 
in  the  United  States,  this  primitive  subfamily  is  very  well  repre- 
sented here  and  contains  some  of  the  largest  and  commonest  ants. 
Notwithstanding  their  superior  size,  in  many  cases  they  cannot  cope 
against  quite  small,  well-armed  ants.  This  was  noted  in  the  genus 
Diacamma,  and  upon  an  invasion  by  such  small  ants,  the  former 
will  grab  bag  and  baggage,  and,  hurrying  out  of  their  nest,  com- 
monly situated  in  a  tree  trunk,  await  the  departure  of  the  marauders. 
Besides,  as  is  generally  the  case  with  the  Ponerinse,  their  colonies 
are  small,  and  sting  and  jaws  do  not  count  for  much  against  a 
superior  number  of  small  ants,  especially  when  many  of  the  latter 
discharge  a  very  disconcerting  fluid  in  their  battles. 

The  Dorylinse  include  the  famous  driver  or  legionary  ants  of  the 
American  and  African  tropics,  that  in  their  foraging  marches  clear 
the  path  of  insects  and  other  creatures.  In  the  Philippines  I  have 
observed  no  such  formidable  ants,  and  the  only  representatives  of 
this  subfamily  familiar  to  me  are  rather  small,  wiry  black  species 
of  the  genus  jEnictus,  that  travel  in  narrow  columns.  The  workers, 
although  blind,  march  with  order  and  great  activity.  They  appear 
to  prey  on  other  ants. 

While  still  on  the  subject  of  ants,  it  would  be  well  to  consider 
briefly  some  habits  of  the  rather  large  muscid  flies,  Bengalia  sp. 
These  are  somber-colored  insects  of  alert  habits  that  hang  around 
the  passing  columns  of  certain  ants,  very  often  those  of  Phidologe- 
ton. It  is  quite  usual  to  see  one  or  more  of  these  flies  perched  right 
near  the  moving  ants,  and  once  in  a  while  to  approach  a  burdened 
ant,  seize  and  snatch  away  its  load  and  to  consume  it  at  a  safe  dis- 
tance. Bengalia,  then,  feeds  upon  the  early  stages  of  ants  and 
whatever  palatable  food  the  ants  may  carry.    Thus,  a  Phidologeton 


332  The  University  Science  Bulletin. 

army  returning,  laden  mostly  with  soft,  white  ants  or  termites,  was 
patronized  by  one  of  these  flies.  The  burdens  being  large,  however, 
necessitated  several  ant-carriers  apiece,  and  thus  made  it  rather 
difficult  for  Bengalia  to  operate. 

Among  the  numerous  arthropods  that  resemble  ants  are  various 
spiders.  That  some  of  these  are  more  or  less  associated  with  the 
ants  they  resemble  is  beyond  doubt,  but  the  whole  subject,  I  be- 
lieve, is  still  in  a  rather  speculative  stage.  There  are  spiders  that  re- 
semble Q^cophylla;  many,  Polyrhachis ;  some,  Diacamma  and  others. 
This  resemblance  is  often  excellent,  though  one  will  learn  quickly 
to  differentiate  ant  and  spider.  The  latter  usually  has  much  the 
better  vision  of  the  two,  and  so  if  disturbed  will  wheel  about  sharply, 
very  unlike  an  ant.  Spiders  have  four  pairs  of  legs,  and  these  ant- 
resembling  species,  in  what  suggests  to  us  an  endeavor  to  mimic 
antennse,  will  wave  the  first  pair  of  legs,  also  unlike  an  ant.  While 
these  spiders  are  slender,  and,  like  ants,  properly  constricted,  the 
large  chelae  do  not  much  resemble  the  jaws  of  ants.  They  have 
somewhat  the  habits  of  attid  spiders. 

SOME  BUTTERFLIES. 

Butterflies  are  not  all  children  of  the  light.  The  tropics  possess 
a  number  of  species  that  are  addicted  to  a  night  life,  or  that  at  least 
avoid  the  sunlight — mainly  somber-colored  insects  that  belong  to 
the  families  Hesperiidae  (or  skippers)  and  Satyridse — and  elsewhere 
they  may  be  seen  at  sunset,  or  perhaps  earlier  on  dark  days,  and 
sometimes  also  before  sunrise,  flying  about,  feeding  or  laying  their 
eggs.  I  have  found  them  coming  to  light  but  rarely,  and  it  seems 
probable  that  they  are  not  so  active  when  the  night  is  far  advanced. 

The  largest  of  these  crepuscular  species  that  I  have  observed  is 
the  coconut  nymphalid,  Amathusia  phidippus  Johanssen,  a  graceful 
brownish  insect  with  a  wing  expanse  of  about  four  inches.  The 
larva  eats  the  leaves  of  the  banana,  the  cocoanut,  and  probably  of 
other  palms. 

The  banana  leaf  roller,  Erionota  thrax  Linne,  is  quite  a  large 
skipper  butterfly,  whose  larva  makes  a  retreat  of  a  strip  of  banana 
leaf,  which  it  cuts  away  from  the  edge  and  more  or  less  parallel  to 
the  midrib  and  rolls  up  as  a  wide  ribbon.  When  it  has  outgrown  or 
consumed  most  of  this  roll,  it  constructs  another  and  larger  one 
and  pupates  in  the  last  one  made.  The  larva  is  covered  with  a 
mealy  white  substance. 

A  skipper  even  larger  than  the  banana  leaf  roller,  and  also  prob- 


Williams:    Philippine  Zoology.  333 

ably  crepuscular,  lays  its  eggs  on  the  rattan  (Calamus),  a  species  of 
thorny  and  climbing  palm.  The  larva  is  covered  with  a  flocculent 
white  material.  It  pupates  in  a  neat  retreat  of  rolled-up  leaflets. 
The  sensitive  pupti  even  when  gently  disturbed,  will  so  move  as  to 
produce  what  is  at  first  a  rather  startling,  whirring  or  scraping  noise. 
Other  large  skippers  feed  also  on  palms,  on  plants  of  the  ginger 
family,  and  on  Araliaceae. 

The  largest  as  ^\'ell  as  one  of  the  handsomest  of  the  day  butter- 
flies is  the  bird-wing  butterfly,  Ornithoptera  nephereus  G.  R.  Gray, 
witli  colors  chiefly  velvety  black  and  brassy  yellow,  and  a  wing  ex- 
panse of  about  six  or  six  and  a  half  inches.  Though  a  fine  insect, 
it  is  by  no  means  the  largest  nor  the  handsomest  of  the  genus.  It  is 
found  in  the  lowlands  to  some  distance  up  in  the  forest.  The  larva 
feeds  on  a  species  of  Aristolochia  or  "Dutchman's  pipe  vine,"  which 
it  shares  with  Papilio  antiphus  Fabricius,  a  much  smaller,  mostly 
black  butterfly.  The  larvae  of  these  two  species,  as  well  as  that  of 
Papilio  philenor  Linn  of  North  America,  resemble  one  another  some- 
what in  that  they  have  fleshy  processes  on  the  body.  The  pupse  also 
are  swollen  at  the  sides  much  more  than  any  other  pupse  of  Papilio 
that  I  know  of. 

The  genus  Ornithoptera  is  sometimes  considered  a  subgenus  of 
Papilio,  and  ranges  south,  well  into  Australia. 

FIREFLIES. 

We  have  fireflies  with  us  practically  throughout  the  year.  This 
is  in  striking  contrast  to  the  comparatively  brief  season  of  the  adult 
beetles  in  the  United  States.  But  there  is  this  much  to  be  said  in 
favor  of  the  latter.  I  have  seen  no  Luzon  island  lampyrid  that 
equals  in  brilliancy  the  light  that  is  emitted  by  Photuris  pennsyl- 
vanica  of  the  Eastern  United  States. 

There  are  quite  a  number  of  species  found  in  this  portion  of  the 
Pliilippines.  Some  have  a  weak  luminosity,  while  others  are  quite 
brilliant,  and  it  is  a  common  thing  to  see  a  whole  bush  or  crown  of 
a  tree  sparkling  like  an  old-fashioned  Christmas  tree,  with  hundreds 
of  these  insects.  Such  trees,  especially  when  isolated,  are  visible 
from  quite  a  distance. 

While  the  larvse  of  fireflies,  being  luminous  themselves,  are  not 
difficult  to  find,  I  do  not  believe  that  many  Philippine  species  have 
yet  been  associated  with  the  adults.  I  have  found  some,  as  related 
by  Fabre  in  Europe,  devouring  snails  within  their  shells,  the  victim 
having  been  overcome  on  some  bush  or  on  the  ground.     In  the 


334  The  University  Science  Bulletin. 

United  States  the  larva  of  the  genus  Phengodes  is  known  to  feed 
upon  myriapods.  Here  at  Los  Bafios  is  a  related  insect  with  a  sim- 
ilar prey.  I  have  kept  some  of  the  larvae  in  captivity  and  fed  them 
living  myriapods  of  at  least  two  species,  and  have  several  times  seen 
the  beetle  larva  overcome  its  large  prey.  It  would  grasp  the  myria- 
pod  by  an  antenna,  and  while  its  victim  might  struggle  violently, 
the  sharp  mandibles  of  the  aggressor  seemed  eventually  to  pierce  the 
antenna  and  to  discharge  through  it  a  quieting  fluid.  At  any  rate, 
the  prey  is  rendered  helpless  without  further  biting,  and  its  interior 
hollowed  out  by  the  voracious  larva. 

OTHER  THAN  INSECTS. 

One  concerned  chiefly  with  the  study  of  insects  cannot,  however, 
fail  to  secure  an  interest  in  other  invertebrates,  as  well  as  verte- 
brates, that  come  to  his  notice. 

The  land  leech  is  one  invertebrate  with  which  those  who  explore 
the  mountain  woods  in  damp  weather  soon  become  acquainted.  If 
the  leech  feeds  solely  on  blood,  it  must  possess  great  powers  of  fast- 
ing. It  is  a  tough,  wormlike  creature,  often  adorned  with  brilliant 
stripes,  and  that  measures  in  the  neighborhood  of  an  inch  in  length. 
It  is  furnished  with  an  anterior  and  posterior  sucker,  by  the  use  of 
which  it  travels  much  after  the  fashion  of  certain  moth  caterpillars 
known  as  inch  worms  or  loopers.  It  is  very  alert  to  a  footfall,  and 
from  some  little  distance,  conscious  of  the  presence  of  man  or  beast. 
Perched  on  an  herb  or  in  the  middle  of  a  trail,  and  raised  up  on  its 
hind  sucker,  it  is  quick  to  grasp  a  passing  leg  with  the  anterior 
sucker  and  climb  on.  Other  leeches  in  the  vicinity  may  w'ave  their 
head  end,  or  move  along  in  haste  in  search  of  this  prospective  meal. 
Once  aboard,  the  hungry  leech  seeks  to  make  a  puncture  in  the 
skin.  This  may  be  a  difficult  matter  in  the  case  of  a  human  being 
whose  legs  are  well  wrapped  in  cloth  puttees,  or  else  very  easy  if 
leather  leggings  or  none  are  concerned.  Firmly  affixed,  it  becomes 
so  filled  with  blood  as  to  present  a  very  rotund  and  inactive  appear- 
ance, and  eventually  to  drop  off.  The  leech,  however,  is  on  the 
whole  much  more  disgusting  than  painful,  as  in  fact  one  may  not 
become  aware  of  its  work  until  it  has  dropped  off,  full  fed.  Like 
the  hobo,  the  leech  does  not  relish  soap,  and  the  barefooted  native 
keeps  this  in  mind  and  secures  partial  protection  by  rubbing  it  on. 

Most  people  look  upon  the  crab  as  a  denizen  of  the  sea,  or  at  least 
as  an  inhabitant  near  the  seashore.  However,  rather  small-sized 
crabs  inhabit  fresh-water  streams  near  Los  Banos,  and  I  have  found 


Williams:   Philippine  Zoology.  335 

them  from  two  to  three  miles  up  such  watercourses,  the  latter  flow- 
ing into  a  large  lake.  One  of  these  crustaceans  was  dug  out  of  a 
decayed  log  along  a  stream  at  an  altitude  of  perhaps  700  feet  on 
Mount  Maquiling. 

Tailless  amphibians  are  numerous  on  Mount  Maquiling,  and  may- 
occur  at  a  considerable  distance  from  permanent  water.  A  quite 
large  species  with  much  of  the  general  appearance  of  our  tree  toads 
patronizes  suitable  vegetation,  and  is  not  averse,  on  occasions,  to 
visiting  houses.  Along  the  streams  are  large  frogs  which  probably 
do  not  equal  our  famous  bullfrog  {Rana  catesbiana)  in  average 
size.  They  are  very  neat  and  fearless  divers,  and  take  surprising 
headers,  from  some  well-chosen  point,  into  the  pool,  many  feet 
below. 

Of  lizards,  skinks  are  the  most  noticeable,  as  they  scurry  away  at 
your  approach  and  rustle  among  the  leaves.  They  are  fond  of  sun- 
ning themselves  along  the  sides  of  paths.  Some  are  over  a  foot  long 
and  rather  stoutly  made.  Very  small  lizards,  presumably  young 
skinks,  are  often  plentiful  in  the  forest.  A  species  of  Draco,  or  fly- 
ing lizard,  seems  to  occur  chiefly  in  the  lower  portions  of  the  forest. 
Its  parachuting  power  is  secured  through  the  extension  of  its  ribs 
beyond  the  body  proper,  so  that  a  wide  sailing  surface  can  be  pro- 
duced. This  expanded  area  is  somewhat  gaudily  colored  and  visible 
only  when  the  lizard  is  in  "flight,"  for  when  at  rest  the  ribs  are 
pressed  alongside  the  body.  Then  the  skinny  and  harmless  little 
creature  shows  nothing  of  its  aeronautic  propensities.  It  is  arboreal, 
and  sails  from  one  tree  to  another  in  an  easy  descent,  making  a 
graceful  upsweeping  landing.  Despite  such  powers,  these  lizards  do 
not  appear  to  wander  far,  as  I  have  seen  them  patronizing  the  same 
tree  for  months. 

Coming  to  grosser  lizards,  the  monitor  lizard  {Varanus  salvator 
Laur.)  is  noticeable  for  its  large  size  and  noisy  haste  when  alarmed. 
It  reaches  a  length  of  several  feet  and  is  pretty  well  at  home  in  the 
water,  on  land  or  up  in  a  tree.  It  no  doubt  consumes  a  variety  of 
food  and  has  quite  a  reputation  as  a  chicken  thief. 

While  snakes  are  common  here,  they  are  not  to  be  found  on  every 
occasion,  and  large  ones  are  somewhat  of  a  rarity.  Pythons  {Python 
reticulatus  Schneid.)  occur  in  this  region,  and  several  years  ago  two 
soldiers  shot  a  specimen  about  twenty-four  feet  long.  Such  exam- 
ples are  few  and  far  between. 

Of  birds,  the  red  jungle  fowl  {Galium  gallus  Linnaeus)  much  like 


336  The  University  Science  Bulletin. 

some  of  our  smaller  domesticated  chickens,  may  be  heard  crowing 
in  the  forest  far  from  human  habitation.  However,  they  not  infre- 
quently associate  with  the  tame  fowl. 

There  are  some  very  gaudy  kingfishers  in  the  woods.  While  some 
favor  streams,  others  may  be  found  in  the  dense  woods  and  must 
be  largely  insectivorous. 

There  are  no  large  carnivorous  animals  here.  Mount  Maquiling 
has  its  full  share  of  wild  pigs,  deer  and  monkeys.  Any  of  these  may 
during  the  night  come  quite  close  to  human  habitations  and  inflict 
minor  damages  to  crops.  Monkeys  travel  in  companies  among  the 
trees  and  feed  upon  the  abundant  fruits.  They  are  not  especially 
noisy  and  their  tails  do  not  appear  at  all  prehensile.  A  powerful 
but  rare  eagle,  Pithecophaga  jefjeryi  Grant,  is  known  to  prey  upon 
monkeys  in  the  Philippines. 


«4 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  12— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  12.) 

ENTOMOLOGY  NUMBER  V 


CONTENTS: 

Notes  on  Nesting  of  Polistes  (Hymenopter.^)  . .  .Dwight  Isely. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIYERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  12. 


Notes  on  Nesting  of  Polistes  (Hymenoptera^  Vespidae).' 

BY  DWIGHT  ISELY, 
Department  of  Entomology,  University  of  Arkansas. - 

BY  FAR  the  most  abundant  paper  wasp  in  northwest  Arkansas 
is  Polistes  metricus  Say.^  Its  nests  can  be  found  almost  any- 
where attached  to  trees  or  shrubbery  or  under  the  eaves  of  buildings. 
They  are  often  quite  numerous.  As,  for  instance,  along  a  stream  for 
a  distance  of  about  three  hundred  yards  the  writer  counted  twenty- 
two  nests  early  in  May,  1921.  In  a  small  block  of  apple  trees, 
somewhat  less  than  an  acre  in  extent,  seven  nests  of  this  species  were 
noted  in  October  of  the  same  year.  About  premises  of  the  insectary 
of  the  United  States  Bureau  of  Entomology  at  Benton ville,  Ark.,  the 
same  season,  five  nests  were  built. 

The  following  notes  relate  largely  to  one  of  these  nests  which  was 
built  just  outside  of  the  screen,  and  under  a  shutter,  of  the  insectary, 
conveniently  situated  for  observation  from  within,  where  complete 
immunity  from  stings  could  be  enjoyed.  The  history  of  the  nest  is 
as  follows: 

The  stem  of  the  nest  was  begun  May  18  by  a  female  wasp.  On 
May  20  three  cells  were  started,  and  three  more  were  begun  the  next 
day.  As  soon  as  the  base  of  each,  of  these  cells  was  built,  an  egg  was 
deposited  in  it;  thus  six  eggs  were  deposited  in  two  days.  From  now 
on  cells  were  added  less  frequently,  but  considerable  time  was  given 
to  enlarging  those  already  started.  On  May  23  two  more  cells  were 
started,  and  the  walls  of  one  of  the  first  cells  were  8  mm.  long.  Two 
more  cells  were  added  May  28.  An  egg  was  deposited  in  one  of 
these,  but  not  in  the  other  until  May  30.    This  marks  the  end  of  the 

1.  Published  with  permission  of  the  Secretary  of  Agriculture. 

2.  Formerly  with  U.  S.  Bureau  of  Entomology:    Deciduous  Fruit  Insect  Investigation. 

3.  Determined  bj-  Mr.  S.  A.  Rohwer,  U.  S.  Bureau  of  Entomology-. 

(341) 


342  The  University  Science  Bulletin. 

first  stage  in  the  history  of  this  nest.    No  more  cells  were  added  un- 
til near  the  time  of  maturity  of  the  offspring. 

Hatching  began  May  30  with  the  three  eggs  which  had  been  de- 
posited on  May  20.  Two  eggs  deposited  on  May  21  hatched  on  May 
31  and  one  on  June  1.  The  incubation  period  of  five  eggs  was  ten 
days  and  of  one  eleven  days.  Records  of  incubation  were  not  com- 
plete for  the  other  eggs,  but  they  required  about  the  same  time. 

The  wasp  larvae  grew  rapidly,  but  at  an  unequal  rate.  This  was 
apparently  due  to  the  fact  that  some  were  favored  in  feeding.  The 
first  grub  spun  the  cap,  closing  its  cell  for  pupation  June  9,  and  the 
other  cells  of  the  first-hatched  larvae  were  closed  June  1,  13,  15,  17 
and  21,  and  July  9,  making  the  larval  period  vary  from  ten  to 
thirty-eight  days.  In  contrast  to  this  extreme  variation,  the  period 
required  for  pupation  was  exactly  eighteen  days  in  all  instances,  the 
wasps  maturing  on  June  27,  July  1,  3,  5,  9  and  27.    All  were  females. 

For  two  days  after  emergence  from  its  cell  the  first  wasp  stayed 
on  the  nest.  After  that  it  began  going  to  the  field,  and  soon  was 
doing  most  of  the  field  work;  that  is,  bringing  in  paper  for  nest 
building  and  partially  crushed  insects  for  larval  food.  Caterpillars 
were  apparently  the  most  frequent  prey.  The  parent  wasp  stayed 
at  home,  received  supplies  from  the  worker,  and  at  first  did  most  of 
the  feeding  of  the  larvae  and  actual  adding  to  the  nest.  As  the  num- 
ber of  offspring  increased,  there  were  always  several  on  the  nest 
ready  to  meet  a  field  worker  and  unburden  it  of  its  supplies.  The 
field  workers  also  often  engaged  in  feeding  the  grubs. 

Shortly  before  the  emergence  of  her  first  offspring,  on  June  21, 
the  parent  wasp  began  three  new  cells  and  deposited  eggs  in  them. 
Three  more  cells  were  started  on  June  27,  and  from  then  on  building 
progressed  rapidly.  By  July  10  there  was  a  total  of  thirty-one  cells 
in  the  nest,  and  all  of  the  cells  vacated  by  maturing  wasps  were 
renovated  and  eggs  were  again  placed  in  them.  At  the  close  of  the 
season  a  total  of  eighty-two  cells  had  been  built. 

This  nest  seemed  to  be  fairly  representative  for  rate  of  growth 
and  size.  Two  other  nests  which  were  started  about  the  same  time 
were  observed  occasionally.  On  July  9  one  had  three  wasps  and 
fourteen  cells;  the  other  had  six  wasps  and  twenty-four  cells.  At 
the  end  of  the  season  they  had  seventy-four  and  102  cells,  respec- 
tively. The  total  number  of  wasps  in  any  of  the  colonies  was  not 
definitely  known. 

The  first  male  wasp  appeared  August  12,  and  after  this  for  at 
least  three  weeks  there  was  a  large  emergence  of  drones.    The  time 


Isely:   Nesting  of  Polistes.  343 

of  appearance  of  the  first  sexually  mature  females  was  not  noted. 
The  males  did  not  go  afield  with  the  regularity  of  the  females,  but 
spent  most  of  their  time  on  the  nest,  so  that  they  made  up  the 
greater  proportion  of  wasps  on  the  nest  during  the  day.  These 
males  were  apparently  waiting  on  the  nest  for  the  emergence  of 
sexually  developed  females.  What  was  probably  a  preliminary  to 
mating  was  observed  twice.  As  a  female  wasp  was  emerging  from 
her  cell  she  was  pounced  on  by  a  male  and  then  by  all  the  males  on 
the  nest.  This  mass  of  wasps  fell  in  a  ball  to  the  ground.  When 
disturbed  by  the  writer,  they  separated.  Mating  was  not  observed. 
A  similar  performance  was  noted  at  another  nest  later  in  the  month. 

Observations  were  not  made  regularly  after  this  time,  the  writer 
having  left  Bentonville,  but  by  the  middle  of  October  all  males  had 
left  the  nest.  A  large  number  of  females  still  collected  there,  but  all 
the  brood  had  emerged  and  all  activity  had  ceased. 

Considerable  opportunity  was  offered  to  observe  the  workings  of 
the  sense  of  direction,  or  rather  the  apparent  lack  of  any  such  sense. 
When  the  colony  consisted  of  only  a  few  individuals  all  of  the  wasps 
apparently  depended  on  local  observations  for  finding  their  way  to 
their  nest.  Any  change  in  the  insectary  shutters  was  confusing  to 
them.  The  oldest  of  the  workers,  when  returning  from  the  field,  was 
observed  to  alight  on  the  shutter  about  a  foot  below  and  two  feet  to 
the  north  of  the  nest.  She  would  run  on  a  horizontal  line  below  the 
nest  and  continue  until  she  was  about  six  inches  past  it,  when  she 
would  turn  at  a  sharp  angle  and  go  directly  to  the  nest.  This  path 
was  always  followed  unless  the  shutter  was  tampered  with,  when 
the  wasp  could  scarcely  find  the  nest  at  all.  None  of  the  other 
wasps  followed  this  same  path,  but  each  apparently  had  its  own 
system. 

The  overwintering  queens  of  Polistes  are  sometimes  gregarious,  a 
number  of  them  starting  a  colony  together  in  the  spring.  I  have 
never  observed  them  working  together  in  building  an  absolutely 
new  nest,  but  on  several  occasions  I  have  seen  a  small  number — 
never  more  than  seven — of  overwintering  queens  renovate  a  large 
nest  left  from  the  season  before  and  start  a  colony  together.  This 
probably  accounts  for  the  very  large  nests  of  several  hundred  cells 
that  frequently  are  found.  These  exceedingly'  numerous  colonies 
were  much  more  pugnacious  than  the  small  colonies.  In  fact,  colo- 
nies of  but  a  few  wasps  are  inclined  to  be  shy  rather  than  pugna- 
cious. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  13— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  13.) 

ENTOMOLOGY  NUMBER  V 


CONTENTS: 

Fi\'E  New  Species  Belonging  to  Genus  Harmolita  (Hy- 
menoptera) W.  J.  Phillips  and  Fred  W.  Poos. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE,  KAN. 


Entered  at  the  post  office  at  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE   BULLETIN 

Vol.  XI v.]  October,  1922.  [No.  13. 


Five  New  Species  Belonging  to  the  Genus  Harmolita 

Motschulsky. 

{Isosoma  Walker  et  Auct.) 

BY  W.  J.  PHILLIPS  AND  F.  W.  POOS, 
L'nited  States  Bureau  of  Entomology,  Charlottesville,  Ya. 

THREE  of  the  five  species  described  in  this  paper  (viz.,  swezeyi, 
panici  and  phalaridis)  would  fall  within  the  genus  Harmolita 
as  restricted  by  Phillips  and  Emery.-  Grahan^  has  more  recently  re- 
defined the  genus,  however,  and  all  five  of  the  new  species  plainly 
fall  within  the  limits  as  he  has  defined  them. 

At  the  time  this  genus  was  revised  by  Phillips  and  Emery,  the 
senior  author  had  seen  only  a  few  specimens  of  H.  cinnce  (n.  sp.  de- 
scribed herewith),  and  since  the  species  differed  somewhat  from  the 
usual  type  of  Harmolita,  he  did  not  consider  it  advisable  to  broaden 
the  scope  of  the  genus  to  include  this  species.  Since  that  time  sev- 
eral similar  species  have  come  under  observation  and  they  seem  to 
be  more  closely  related  to  Harmolita  than  to  Eurytoma  from  a  bio- 
logical standpoint,  as  well  as  from  the  standpoint  of  external  char- 
acters and  the  character  of  the  ovipositor.  The  writers  fully  agree 
with  Gahan  that  it  is  more  advisable  to  include  these  species  in  the 
genus  Harmolita  than  to  erect  a  new  genus  for  their  reception. 

The  principal  characters  assigned  to  the  genus  by  Gahan  are  as 
follows:  Abdomen  of  female  elongate,  conical  or  subfusiform,  with 
segments  more  or  less  subequal,  the  fourth  segment  (when  prop- 
odeum  is  considered  a  thoracic  segment)  never  greatly  enlarged  as 
in  Eurytoma.  Occiput  slightly  concave  and  immargined.  Antennae 
inserted   at  or  above  middle  of   face;   flagellum  weakly   clavate; 

1.  Order  Hymenoptera,  family  Eurytomidae. 

2.  U.  S.  Nat.  Mus.,  vol.  5.'),  pp.  443-471,  plates  39-48. 

3.  Proc.  U.  S.  Nat.  Mus.,  vol  61,  1922,  art.  24,  p.  7. 

(349) 


350  The  University  Science  Bulletin. 

funicle  usually  five- jointed  and  club  three- jointed,  though  in  some 
cases  funicle  is  six-jointed  and  club  two-jointed.  Propodeum  not  or 
scarcely  longei|  than .  scutellum,  not  sharply  declivous,  usually 
rugosely  sculptured;  with  a  more  or  less  distinct  median,  longi- 
tudinal depression.  Sculpturing  of  head  and  dorsum  of  thorax  either 
reticulate  and  shining,  rugulose  punctate  without  umbilicate  punc- 
tures or  rugulose  with  a  few  more  or  less  indefinite  umbilicate  punc- 
tures. 

Prof.  C.  R.  Crosby,  of  Cornell  University,  kindly  placed  his  speci- 
mens and  notes  on  H.  phalaridis  at  the  disposal  of  the  writers.  Pro- 
fessor Crosby  found  that  this  species  has  two  generations  a  year, 
each  generation  confining  itself  to  Phalaris  arundinacea.  The  re- 
maining species  were  reared  from  dried  grasses  collected  in  the  field. 
No  further  observations  on  their  biology  have  thus  far  been  made. 

Acknowledgment  is  made  to  Mr.  A.  B.  Gahan  of  the  United  States 
National  Museum  for  his  kindly  criticism  of  the  manuscript. 

Harmolita  swezeyi,  n.  sp. 

(PI.  XXXVI,  figs.  2  and  5;  pi.  XXXVII,  fig.  7.) 

Female.  Length,  2.90  mm.  Prescutum  reticulately  lineolate  and  without 
broad,  shallow  impressions  or  umbilicate  punctures.  Pronotal  spots  large, 
occupying  about  one-half  of  the  anterior  margin  of  prothorax,  spots  dull 
yellowish.  Sometimes  entire  thorax  and  propodeum  brownish.  Propodeum 
without  a  distinct  margined,  median,  longitudinal  groove,  though  there  is 
usually  a  distince  median  longitudinal  depression.  Propodeum  rugulose  an- 
teriorly and  usually  granulose  in  the  remaining  portion.  Spiracular  carinas 
usually  distinct  but  weak.  Abdomen  equal  to  or  slightly  longer  than  head  and 
thorax  combined;  conically  pointed;  second  segment  equals  one-fourth  length 
of  abdomen;  remaining  segments  subequal,  though  five  is  longest  and  three 
shortest.  Legs  in  black  specimens  are  black  except  at  knees  and  tarsi,  which 
are  luteous;  in  brownish  specimens. the  legs  approach  the  color  of  the  thorax, 
knees  and  tarsi  lighter. 

Antennae:  Funicle  five-jointed;  first  funicle  plus  ring  joints  longer  than 
pedical;  all  of  funicle  joints  usually  distinctly  longer  than  broad;  middle  joint 
of  club  apparently  quadrate;  scape  and  pedical  yellowish. 

Species  medium  to  small.  Median  line  of  face  below  insertion  of  antennae 
without  markings. 

Males.    Unknown. 

This  species  will  run  to  the  couplet  separating  agrostidis  and  websteri  in  the 
Phillips  and  Emery  table  of  species,  but  may  be  readily  distinguished  from 
either  species  by  the  yellowish  scape  and  the  longitudinal  depression  in  the 
propodeum. 

Type  locality.    Honolulu,  Hawaii. 

Type.    Cat.  No.  25,471,  U.  S.  N.  M. 

Described  from  thirteen  females  reared  from  stems  of  Bermuda  grass 
{Cynodon  dactylon),  in  Honolulu,  Hawaii,  by  Mr.  O.  H.  Swezey. 


Phillips  and  Poos:   Harmolita.  351 

Harmolita  panici,  n.  sp. 

(PI.  XXXVI,  figs.  7  and  8;   pi.  XXXVII,  figs.  6  and  8.) 

Female.  Length.  3  mm.  Prcscutvim  nijinlose;  pronotal  spots  small,  occupj'- 
ing  one-half  or  less  of  the  anterior  dorsal  margin  of  the  prothorax ;  spots  dull. 
Propodeum  with  or  without  a  distinct  continuous,  median,  longitudinal  groove, 
though  there  is  a  distinct  indication  of  a  groove  anteriorly;  if  groove  is  con- 
tinuous it  is  faint  and  very  shallow,  and  not  very  clearly  margined ;  rugulose 
within  and  laterad  of  groove.  Abdomen  rather  slender;  longer  than  head  and 
thorax  combined;  second  segment  comprising  one-fourth  to  one-fifth  length  of 
abdomen  exclusive  of  1 ;  3  shortest;  4,  5  and  6  approximately  the  same  length. 
Legs  black  except  tarsi,  knees  and  the  lower  face  of  the  front  femora,  which 
are  luteous. 

Antennje:  Funicle  five-jointed;  first  segment  plus  ring  joint  slightly  longer 
than  pedicel ;  first  and  second  segment  about  equal  in  length ;  3,  4  and  5  about 
equal  in  length,  but  each  shorter  than  either  1  or  2;  all  segments  longer  than 
broad.  Antennae  black ;  median  line  of  face  below  insertion  of  antennse  without 
markings. 

Species  small  to  medium. 

Male.  Length,  2.51  mm.  Prescutum  as  in  female.  Pronotal  spots  minute, 
scarcely  visible  from  abo\e.  Propodeum  without  a  distinct  median  groove; 
rugulose.  Petiole  granulose,  somewhat  shorter  than  hind  coxse.  Legs  colored 
as  in  female  except  that  front  femora  are  darker. 

Antennse:  Scape  almost  same  width  throughout,  as  seen  in  lateral  profile, 
with  no  distinct  shoulder  near  distal  extremity.  First,  second  and  third 
flagellar  joints  with  two  to  three  annulations  at  distal  extremity;  bristles  nu- 
merous and  short,  scarcely  half  the  length  of  the  segments. 

The  individuals  of  this  species  without  the  propodial  groove  run  to  couplet 
14  in  the  Phillips  and  Emery  table  of  species,  but  can  be  easily  separated  from 
hordei  by  having  the  legs  not  red,  and  from  trilici  by  being  a  smaller  species 
and  the  sculpturing  being  much  smoother.  The  individuals  that  have  the 
propodial  groove  run  to  couplet  18,  in  which  vaginicola  and  secalis  are  sepa- 
rated. Both  of  these  species  are  longer  and  more  coarsely  sculptured;  vagini- 
cola has  yellowish  antennal  scapes  and  the  propodeum  of  secalis  is  granulose, 
which  will  easily  separate  them  from  panici. 

Type  locality.    Charlottesville,  Va. 

Type.    Cat.  No.  25,472,  U.  S.  N.  M. 

Described  from  three  females  and  one  male  reared  from  stems  of  Panicum 
clandestinum  at  Charlottesville  by  the  junior  author. 

Harmolita  phalaridis,  n.  sp. 

(PI.  XXXVI,  figs.  6  and  9;   pi.  XXXVII,  figs.   1  and  2.) 

Female.  Length,  3.70  mm.  Prescutum  reticulately  lineolate  with  numerous 
broad,  shallow  impressions;  very  few  such  impressions  on  pronotum,  but  scu- 
tellum  is  quite  thickly  pitted.  Pronotal  spots  bright  and  large,  occupying 
about  two-thirds  anterior  dorsal  margin  of  prothorax.  Propodeum  with  a  dis- 
tinct, continuous,  median,  longitudinal  groove,  which  is  usually  margined; 
usually  deep  throughout,  though  often  shallow  posteriorly;  numerous  cross 
rugae  within  groove,  but  no  indication  of  central  carina;  very  rugulose  laterad 
of  groove;  spiracular  carinas  prominent  and  spiracular  area  usually  well  de- 


352  The  University  Science  Bulletin. 

fined.  Abdomen  same  length  as  head  and  thorax  combined;  segments  3  and  5 
about  of  equal  length;  4,  6  and  7  about  equal  in  length  and  each  usually  longer 
than  either  3  or  5.  Legs:  All  knees,  tibiae  and  tarsi  usually  reddish  brown, 
femora  blackish. 

Antennae:  Funicle  five-jointed;  first  funicle  joint  plus  ring  joint  about  twice 
as  long  as  pedicel;  first  joint  of  funicle  slender  and  same  size  throughout,  the 
distal  tip  somewhat  flaring;  all  of  funicle  joints  distinctly  longer  than  broad; 
club  joints  also  longer  than  broad;  antennae  black  and  \ery  slender. 

Species  medium  in  size. 

Male.  Length  2.60  mm.  Prescutum  as  in  female,  but  there  are  few  thor- 
acic punctures;  pronotal  spots  large  and  bright.  Propodeum  with  or  without 
a  groove;  groove  when  present  is  often  rather  poorly  defined;  propodeum 
usually  very  rugose,  though  it  may  sometimes  be  granulose;  spiracular  area 
usually  well  defined.  Petiole  usually  about  twice  as  long  as  broad,  granulose 
and  extends  beyond  the  tip  of  the  coxae.  Legs:  All  knees  and  tarsi  testaceous; 
front  tibiae  usually  reddish  brown. 

Antennae :  Flagellum  with  peticel  longer  than  head  and  thorax  combined ; 
hairs  on  first  flagellar  joint  approximate! j'  same  length  as  those  on  last  joint; 
last  joint  bears  a  slender  tubercle  at  end  about  twice  as  long  as  broad.  Scape, 
exclusive  of  base,  a  little  o\'er  twice  as  long  as  broad,  broadest  about  center, 
as  seen  in  lateral  profile ;  scape  as  seen  in  lateral  profile  nearly  twice  as  broad 
as  first  flagellar  joint.  There  are  four  or  more  annulations  at  each  articulation 
of  the  flagellum. 

This  species  iims  to  dactylicola  in  the  Phillips-  and  Emery  table  of  species, 
but  the  females  may  be  separated  by  the  following  characters:  H.  phalaridis 
has  more  densely  pitted  scutellum ;  propodeum  more  rugulose ;  groove  deeper 
and  same  width  throughout;  spiracular  carinse  more  prominent;  tibiae  usually 
reddish  brown;  first  funicle  joint  of  antennae  cylindrical,  very  slender  and, 
distal  extremity  somewhat  flaring  at  tip.  All  segments  of  antennae  more  slen- 
der than  in  dactylicola. 

Type  locality.    Ithaca,  N.  Y. 

Type.    Cat.  No.  25,473,  U.  S.  N.  M. 

Described  from  manj^  males  and  females  reared  from  stems  of  Phalaris 
arundinacea  collected  at  Ithaca,  N.  Y.,  by  Professor  Crosby  and  the  junior  au- 
thor, and  from  specimens  reared  from  stems  of  Phalaris  sp.  collected  at  Elk 
Point,  S.  Dak.,  by  Mr.  C.  N.  Ainslie  of  the  U.  S.  Bureau  of  Entomology. 

Harmolita  cinme,  n.  sp. 

(PI.  XXXVI,  figs.  1  and  3;   pi.  XXXVII,  figs.  3  and  4.) 

Female.  Length,  3.80  mm.  The  whole  thorax  somewhat  rugulose  and  more 
or  less  distinctly  umbilicately  punctured,  the  pimctures  shallow  and  usually 
not  well  defined;  prescutum  sometimes  not  umbilicately  punctured  in  anterior 
third;  pronotal  spots  small,  occupying  about  one-third  anterior  dorsal  margin 
of  pronotum,  visible  from  above.  Proi)odeum  with  a  deep,  margined,  con- 
tinuous, median,  longitudinal  groove  of  medium  width;  groove  with  numer- 
ous cross  rugae  and  usually  with  a  central  longitudinal  carina;  very  rugulose 
laterad  of  groove;  sjiiracular  area  usually  well  defined,  thovigh  sometimes  the 
spiracular  carinae  are  weak.  Abdomen  equal  to  or  slightly  longer  than  head 
and  thorax  combined,  and  almost  as  pointed  as  the  average  Harmolita;  seg- 


Phillips  and  Poos:    Harmolita.  353 

ment  2  occupies  between  one-third  and  one-fourth  the  length  of  abdomen; 
segments  3,  4,  5,  6  and  7  approximately  equal  in  length.  Legs  often  variable 
in  color;  sometimes  the  legs  are  black  throughout  except  the  knees,  front 
tibiae  and  all  tarsi,  which  are  luteous;  perhaps  more  often  the  basal  third  to 
half  of  front  and  middle  femora  and  basal  two-thirds  of  hind  femora  blackish; 
all  tibiae,  knees  and  tarsi  almost  reddish  brown. 

Antennae:  Funicle  five-jointed;  club  three-jointed;  first  funicle  joint  plus 
ring  jointly  nearly  twice  as  long  as  pedicel;  all  segments  longer  than  broad;  the 
first  two  funicle  joints  longest,  the  remaining  ones  of  approximately  the  same 
length.  Antennae  black.  Median  line  of  face  below  insertion  of  antennae 
slightl}'  elevated;  doreally  it  appears  almost  carinate. 

Species  medium  to  large. 

Male.  Length,  2.90  mm.  Sculpturing  of  thorax  very  much  the  same  as  in 
female,  except  that  the  umbilicate  pimctures  are  not  nearly  so  distinct;  pro- 
notal  spots  verj'  small,  often  scarcely  visible  from  above.  Propodeum  variable; 
there  may  be  a  deep,  rather  broad,  margined,  median  longitudinal  groove, 
very  rugulose  within  and  laterad  of  groove,  or  the  groove  may  not  be  con- 
tinuous and  shallow,  and  it  may  be  granulose  within  and  laterad  of  the  groove. 
In  the  latter  case  the  petiole  is  usually  granulose;  when  the  propodeum  is 
very  rugulose  the  petiole  is  usuall}/^  somewhat  rugulose.  Petiole  long,  slender; 
the  tip  of  the  hind  coxae  often  extending  only  to  about  the  middle  of  petiole. 
Legs  colored  as  in  female. 

Antennae:  Longer  than  head  and  thorax  combined;  scape  as  seen  in  profile 
thickened  somewhat  at  center;  no  distinct  club;  first  segment  of  flagellum 
approximately  as  long  as  scape;  the  remaining  segments  about  of  equal  length; 
segments  excised  with  about  three  aimulations  at  the  distal  extremity  of  all 
except  distal  segment. 

Type  locality.    Youngstown,  Ohio. 

Type.    Cat.  No\  25,474,  U.  S.  N.  M. 

Described  from  many  females  and  eight  males  reared  from  stems  of  Cinna 
arundinacece  collected  at  Youngstown,  Ohio,  by  Mr.  W.  T.  Emery  and  at  Niles, 
Ohio,  by  the  junior  author. 

Harmolita  phalaricola,  n.  sp. 

(PI.  XXXVI,  figs.  4  and  10;  pi.  XXXVII,  fig.  5.) 

Female.  Length,  3.52  mm.  Prescutum  somewhat  rugulose,  and  the  whole 
thorax  bearing  numerous  but  rather  indefinite  umbilicate  punctures.  Pronotal 
spots  dull,  minute,  scarcely  visible  from  above. 

Propodeum  with  a  distinct,  continuous,  deep,  medium  to  narrow  longitudinal 
median  groove;  groove  not  distinctly  margined  throughout;  very  rugulose 
within  and  laterad  of  groove;  spiracular  area  not  clearly  defined  by  spiracular 
carinas.  Abdomen  short  and  thick,  approaching  ovate;  slightly  shorter  than 
head  and  thorax  combined;  segment  2  occupying  between  one-third  and  one- 
half  dorsal  length  of  abdomen;  segments  vary  in  length  as  is  common  in 
Harmolita,  due  to  telescoping  of  segments  when  the  insects  die ;  3  and  4  often 
nearly  same  length;  5,  6,  and  7  often  about  same  length,  but  shorter  than 
either  3  or  4.  Legs:  Basal  half  of  front  and  basal  two-thirds  of  middle  and 
hind  femora  black ;  remaining  portion  of  legs  usually  reddish  brown. 


354  The  University  Science  Bulletin, 

Antennae:  Funicle  apparently  six-jointed  and  club  two-jointed;  first  funicle 
plus  ring  joint  about  twice  the  length  of  the  pedicel;  segments  4,  5  and  6  about 
quadrate;  club  joints  nearly  quadrate  also. 

Species  medium  to  large. 

Males.    Unknown. 

Type  locality.    Elk  Point,  South  Dakota. 

Type.    Cat.  No.  25,475,  U.  S.  N.  M. 

Described  from  ten  females  reared  from  stems  of  Phalaris  sp.  collected  at 
Elk  Point,  S.  Dak.,  by  C.  N.  Ainslie  of  the  United  States  Bureau  of  Ento- 
mology. 


356  The  University  Science  Bulletin. 


PLATE  XXXVI. 

1.  Ovipositor  of  H.  cinnce. 

2.  Ovipositor  of  H.  swezeyi. 

3.  Propodeum  of  H.  cinnce. 

4.  Propodeum  of  H.  phalaricola. 

5.  Propodeum  of  H.  swezeyi. 

6.  Propodeum  of  H.  phalaridia. 

7.  Propodeum  of  H.  panici. 

8.  Ovipositor  of  H.  panici. 

9.  Ovipositor  of  H.  phalaridis. 
10.    Ovipositor  of  H.  phalaricola. 


Phillips  and  Poos:    Harmolita- 


357 


PLATE  XXXVI. 


:S!* 


358  The  University  Science  Bulletin. 


PLATE  XXXVII. 

1.  Antenna  of  the  male  of  H.  phalaridis. 

2.  Antenna  of  the  female  of  H.  phalaridis. 

3.  Antenna  of  the  male  of  H.  cinnoe. 

4.  Antenna  of  the  female  of  //.  cinnoe. 

5.  Antenna  of  the  female  of  H.  phalaricola. 

6.  Antenna  of  the  male  of  H.  panici. 

7.  Antenna  of  the  female  of  H.  swezeyi. 

8.  Antenna  of  the  female  of  H.  panici. 


Phillips  and  Poos:    Harmolita. 


359 


PLATE  XXXVII. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  14— October,  1922. 

(Whole  Series,  \'ol.  XXIV,  No.  14.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

The  Urinary  System  of  Phlegethontius  sexta 

(Lepidoptera) G.  H.  Vansell. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post,  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  14. 


The  Urinary  System  of  Phlegethontius  sexta  Johan. 

(Lepidoptera) . 

BY  G.  H.  VANSELL. 

(Contribution  from  the  Zoological  Laboratory  of  the  University  of  Kentucky.) 

PHLEGETHONTIUS  SEXTA  Johan.,  commonly  known  as  the 
southern  tobacco  worm,  is  used  in  many  laboratories  for  mor- 
phological study  material.  The  paper  of  Alvah  Peterson,  published 
in  the  September  number  of  the  "Annals  of  the  Entomological  So- 
ciety of  America,  1912,"  treats  the  general  anatomy  of  Protoparce 
Carolina  Linn.,  but  the  Malpighian  tubules  are  not  shown  in  his  fig- 
ures as  fully  as  some  other  parts.  If  the  conditions  shown  in  the 
figures  accompanying  this  paper  differ  from  those  found  by  Peterson 
in  his  work  it  may  throw  some  light  on  the  proposed  synonomy  of 
the  species  in  question. 

Malpighian  tubules  are  usually  two  in  number,  or  in  multiples  of 
two,  and  in  most  cases  they  empty  directly,  or  through  a  bladder, 
into  the  intestine.  In  the  larvae  of  Microgaster,  Koulagin  found  that 
they  open  dorsally  on  the  outside  of  the  body  on  each  side  of  the 
anus.  Those  of  Phlegethontius  sexta  Johan.  empty  into  the  small 
intestine. 

.  The  specimens  used  for  dissection  in  this  work  were  killed  in  va- 
rious ways.  Some  were  dropped  into  a  solution  of  50  per  cent  alco- 
hol. 48  per  cent  water,  and  2  per  cent  formalin.  Immediately  after 
death  they  were  removed  and  the  body  wall  split  to  allow  the  free 
passage  of  the  preservative  to  the  inside  organs.  At  the  time  of  use 
these  specimens  were  further  hardened  by  adding  a  solution  of  picric 
acid  and  chloral  hydrate.  Others  were  dropped  into  Bouin's  picro- 
formol  mixture,  and  later  run  up  through  alcohols.    Various  stains 

(36.5) 


366  The  University  Science  Bulletin. 

were  used  upon  that  material  which  was  imbedded  for  sectioning. 
Borax  carmine  and  Delafield's  hemotoxylin  seemed  to  give  the  best 
results. 

The  Malpighian  tubules  of  Phlegethontius  sexta  are  six  in  number 
and  of  a  light  yellowish  color.  Four  of  these  are  located  largely 
laterodorsally  to  the  intestinal  tract,  the  other  two  being  ventrally 
situated.  Each  tubule  is  apparently  free  at  the  end  distad  from  the 
bladder.  There  are  two  bladders,  located  one  on  each  side  between 
the  ventriculus  and  the  small  intestine  (fig.  2),  and  just  posterior  to 
the  ventriculus  they  empty  separately  into  the  small  intestine 
through  short,  smooth  tubes  (fig.  3).  From  the  opposite  end  of 
either  bladder  a  tube  arises  which  runs,  dorsally  and  branches  im- 
mediately into  two;  one  of  these  tubes  runs  ventrally  and  forward, 
the  other  dorsally,  and  this  one  divides  into  two  more  as  it  nears 
the  top  of  the  ventriculus.  These  six  tubes  run  forward  toward  the 
head  and  turn  back  posteriorly  at  about  the  second  abdominal  seg- 
ment, as  shown  in  figures  5  and  6.  Posterior  to  the  bladders  all 
these  tubes  interweave  with  the  adipose  tissue  until  they  lose  their 
identity. 

On  high  magnification  the  tubes  present  an  interesting  appear- 
ance, in  that  each  one  is  covered  with  small  nodules.  These  nodules 
become  more  numerous  on  the  tubes  the  greater  the  distance  from 
the  connection  with  the  bladder,  until  at  the  distal  ends  the  nodules 
are  very  closely  crowded  together  (figs.  7  and  8).  Tracheae  run  to 
the  tubes  and  branch  into  invisible  threads  upon  the  surface  of  the 
nodules  (fig.  9). 

Both  the  nodules  and  the  tubes  are  hollow,  and  an  opening  from 
each  nodule  leads  into  the  tube,  making  a  continuous  cavity  to  all 
parts  (figs.  10,  11  and  12).  The  exact  histological  nature  of  the 
tubes  is  very  hard  to  determine  on  account  of  their  delicate  nature. 
The  figures  showing  these  structures  are  not  shown  here,  for  more 
work  is  being  done  upon  them  and  a  later  paper  will  appear. 

LITERATURE. 

Berlese,  Antonio.  1909.  Gli  Inset ti  lore  organizzazione,  sviluppo,  abitudini 
e  rapporti  coll'uomo,  pp.  779-788;  figs.  971,  972,  975.  Societa  Editrice  Li- 
braria,  Milano. 

FoLSOM,  J.  W.  1906.  Entomology,  with  Special  Reference  to  its  Biological 
and  Economic  Aspects,  pp.  123-124.    Philadelphia;  P.  Blakistons  &  Co. 

Peterson,  Alvah.  1912.  Anatomy  of  the  Tomato  Larva,  Protoparce  Carolina. 
Annals  of  the  Entomological  Society  of  America,  pp.  245-272;  pis.  XIX- 
XXI.    Columbus,  O. 


368  The  University  Science  Bulletin. 


PLATE  XXXVIII. 

Fig.  1.  Dorsal  view  of  the  region  into  which  the  Malpighian  tubules  empty. 
bl,  bladder;  dd,  Dorsal  duct;  ve,  ventriculus;  si,  small  intestine. 

Fig.  2.  Ventral  view  of  the  same  region,  bl,  bladder;  cd,  common  bladder 
duct;  vd,  ventral  duct;  dd,  dorsal  duct;  n,  nodule  on  the  Malpighian  tubule; 
si,  small  intestine. 

Figs.  3,  4.  Lateral  view,  showing  the  left  and  right  sides  of  the  intestine 
with  the  bladders  and  ducts,  ve,  ventriculus;  si,  small  intestine;  bl,  bladder; 
o,  duct  opening  into  the  small  intestine. 

Fig.  5.  Dorsal  view  of  the  ventriculus,  showing  the  arrangement  of  the 
Malpighian  tubules.  2nd,  second  abdominal  segment;  9th,  ninth  abdominal 
segment;  xx,  region  in  which  the  free  ends  of  the  Malpighian  tubules  and 
the  adipose  tissue  interweave  into  a  dense  mass. 

Fig.  6.  Ventral  view  of  the  ventriculus,  showing  the  arrangement  of  the 
Malpighian  tubules.    Labels  as  in  figure  5. 

Fig.  7.  Malpighian  tubules  under  the  microscojie,  showing  the  nodules. 
This  piece  of  the  tube  occurred  just  posterior  to  where  a  dorsal  tube  turns 
caudad.    t,  tubule;  n,  nodule. 

Fig.  S.  Section  from  the  Malpighian  tubule  toward  the  distad  end,  just 
anterior  to  the  highly  convoluted  area.  The  nodules  are  quite  numerous  here 
and  the  tube  thickened,    t,  tube  n,  nodule. 

Fig.  9.  Highly  magnified  portion  of  the  tubule,  showing  the  disappearance 
of  the  tracl{ae  on  the  nodules,    n,  nodule;  t,  tube;  tr.  trachea'. 

Fig.  10.    Cross  section  of  tubule. 

Fig.  n.    Cross  section  of  tubule  and  nodules. 

Fig.  12.   Longitudinal  section  of  tubule  and  nodule. 


Vansell:    Phlegethontius  Sexta. 


369 


PLATE  XXXVIII. 


16 


11 


12 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  15— October,  1922. 

(Whole  Series,  Vol.   XXIV,  No.   15.) 

ENTOMOLOGY  NUMBER  V 


CONTENTS: 

A  Brief  Re.sume  of  Investig.ations  Made  in   1913  on 

Trogoderm.a  inclus.\  (Coleoptera) Adolph  H.  Beyer. 


PUBLISHED  BY  THE  UNIVERSITY 

LAWRENCE.  KAX. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIYERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  15. 


A  Brief  Resume  of  Investigations  Made  in  1913  on 
Trogoderma  inclusa  Lee.   (a  Dermestid) . 

By  ADOLPH  H.  BEYER. 

THE  work  herewith  reported  was  undertaken  at  the  suggestion 
of  Prof.  S.  J.  Hunter,  who  had  received  complaints  concerning 
damage  to  leather  horse  collars.  The  damage  consisted  in  more  or 
less  numerous  perforations  of  the  leather  by  some  insect,  which 
injury  interfered  seriously  with  the  marketability  of  the  manu- 
factured goods.  It  became  my  task  to  determine  the  insect  causing 
the  damage,  the  source  of  infestation,  and  the  methods  of  con- 
trolling the  pest.  The  investigations  were  made  in  1913,  but  are 
reported  here  for  the  first  time. 

NATURE  OF  THE  INJURY  TO  THE  HORSE  COLLARS. 

The  leather  of  the  infested  collars  has  the  appearance  of  being 
perforated  with  numerous  little  round  holes  about  the  size  of  a  pin- 
head.  The  insects  reared  from  infested  collars  proved  to  be  der- 
mestid beetles  belonging  to  the  above-named  species.  The  insects 
hatch  in  the  rye  straw  used  in  filling  the  collars,  and  the  larvae  live 
upon  the  grain  left  in  the  straw.  As  the  larvse  develop  to  within  a 
month  of  the  adult  stage,  they  approach  the  outside  of  the  collar 
and  emerge  by  eating  holes  through  the  leather  of  the  collar. 

A  number  of  firms  engaged  in  the  manufacture  of  collars  have 
reported  damage  of  this  character.  One  firm  in  Texas  writes: 
"We  wish  to  state  that  five  or  six  years  ago  we  had  trouble  of  this 
kind  owing  to  the  fact  that  we  were  using  rye  straw  with  lots  of 
grain  still  in  the  straw,  and  small  worms  or  insects  would  bore 
through  the  leather  into  the  collar  to  get  at  the  grain.  We  had  very 
little  loss,  as  we  soon  found  out  the  trouble,  and  therefore  quit 
using  straw  that  had  any  grain  in  it  at  all." 

(373) 


374  The  University  Science  Bulletin. 

Two  or  three  firms  securing  straw  from  a  grower  near  Lawrence, 
Kan.,  reported  serious  trouble.  An  examination  of  one  factory  re- 
vealed fifty  per  cent  of  the  collars  infested.  Upon  examining  the 
floor,  walls  and  window  and  door  casings,  the  cracks  were  found  to 
contain  many  adult  beetles,  which  in  all  cases  were  dead.  The 
light  and  heat  of  the  sun  had  a  marked  influence  upon  the  distribu- 
tion and  development  of  the  insects,  as  the  collars  near  the  south 
windows  were  more  seriously  damaged,  the  injury  decreasing 
gradually  as  one  neared  the  north  side  of  the  building. 

SOURCE  OF  INFESTATION. 

Since  injury  was  reported  from  manufacturers  using  straw  grown 
near  Lawrence,  Kan.,  it  appeared  possible  that  the  straw  might  have 
been  infested  before  it  was  shipped  to  the  factories.  An  investiga- 
tion was  made  first  at  the  ranch  where  the  rye  straw  was  grown. 
It  was  found  that  the  sheaves  were  stacked  up  in  the  barns  to  con- 
serve the  value  of  the  straw  for  collar-stuffing  purposes,  and  during 
the  winter  months  a  specially  made  thresher  was  used  in  removing 
the  grain  from  the  head  of  the  sheaf  without  cutting  the  band.  After 
the  rye  bundles  were  threshed  they  were  baled,  being  compressed 
and  tied  with  wire  ready  for  shipping  to  the  collar  factory.  I  found 
no  trace  of  the  insect  about  any  part  of  the  sheaves.  They  were 
apparently  free  in  every  respect  from  the  insect.  I  also  examined 
the  threshed  bundles  before  they  were  baled,  and  noticed  that  there 
was  considerable  grain  left  in  the  straw,  which  later  afforded  food 
for  the  development  of  the  insects  in  the  collar.  I  next  examined 
the  threshed  rye  in  the  bin,  thinking  perhaps  the  insects  might  be 
hibernating  at  this  season  of  the  year.  I  collected  several  ciuarts  of 
the  rye  in  a  screw-sealed  glass  jar,  to  keep  out  other  infestations, 
and  took  this,  with  one  of  the  rye  sheaves,  to  the  laboratory. 

I  placed  some  of  the  rye  straw  taken  from  the  sheaf  in  a  large 
glass-enclosed  cylinder.  This  was  placed  in  the  incubator  at  ordi- 
nary room  temperature  of  70°  F.  I  recorded  the  temperature  each 
day  and  watched  for  the  development  of  infestations,  but  noted  no 
development  of  life  for  several  weeks.  At  the  end  of  three  weeks 
I  found  two  species  of  grain  weevil  and  several  parasites,  which  I 
removed  from  the  jar  and  placed  in  separate  vials.  At  the  end  of  the 
fourth  week  I  removed  from  the  jar  five  specimens  of  larva.^,  ap- 
parently of  the  same  species  of  Dermestidse  that  was  the  cause 
of  the  collar  infestation.  April  15,  the  adults  emerged  from  the 
pupa^  cases.  I  found  them  to  be  the  same  species  as  those  causing 
the  damage  to  the  collars. 


Beyer:    Trogoderma  inclusa.  375 

EXTENT  OF  INFESTATION. 

x\s  stated  before,  fifty  per  cent  of  tlie  colhirs  in  one  factory  were 
damaged.  I  made  a  careful  dissection  of  both  sides  of  a  badly 
infested  collar  by  removing  the  leather  of  the  facing  of  the  collar. 
I  have  found  only  llie  larval  stage  of  the  life  cycle  in  the  collars 
so  far  examined.  T  made  an  approximate  count  of  all  the  larvae 
found  in  this  collar,  and  900  larvae  was  the  result.  I  also  counted 
the  number  of  holes  in  the  same  collar  leather,  to  get  the  per  cent 
of  damage  done  by  each  insect.  The  actual  number  of  holes  in  this 
collar  made  by.  this  pest  was  424.  Approximately  1,324  of  the  in- 
sects infested  this  collar. 

The  larvae  varied  from  2i/^  to  5  mm.  in  length.  They  were 
usually  found  in  the  heads  of  the  rye,  or  eating  on  the  scattered 
grains.  The  larvae  eat  straight  across  the  ends  or  sides  of  the  grain, 
something  as  a  mouse  gnaws,  and  not  after  the  fashion  of  the  grain 
borers.  Wherever  the  tunneling  or  eating  process  was  found,  I  also 
found  evidence  of  molting.  I  found  small  larvae  emerged  in  heads 
of  rye,  which  is  e\'idtnce  that  the  eggs  had  been  laid  in  the  heads. 

LIFE  HISTORY  OF  THE  BEETLE. 

In  making  dissections  of  the  collars  which  had  been  incubated 
at  a  constant  temperature  of  70  degrees  for  several  weeks,  I  found 
various  sizes  of  larvae  and  a  number  in  the  pupal  state.  None 
was  found  in  the  adult  state.  No  eggs  were  found  in  the  collar  dis- 
sections. 

MATING  AND  OVIPOSITION. 

The  beetles  I  found  in  all  cases  mated  a  day  or  two  after  emerging 
from  the  pupal  skins.  The  eggs  varied  in  number  from  ten  to  fifty. 
They  were  placed  around  indiscriminately  on  the  bottoms  of  Petri 
dishes  from  four  to  six  days  after  copulation.  In  a  number  of  cases 
the  female  oviposited  on  the  rye  placed  in  the  dishes,  and  the  eggs 
adhered  to  the  rye  by  means  of  little  filaments  projecting  from  one 
end  of  the  egg.  The  young  larvae  hatched  from  eight  to  twelve 
days  later  at  ordinary  room  temperature.  The  young  larvae,  soon 
after  hatching,  began  to  feed  on  the  material  at  hand.  They  did 
not  wander  unless  food  was  scarce  or  poor.  I  reared  them  entirely 
upon  rye  grain.  The  growth  of  the  larvae  depends  to  a  considerable 
extent  upon  temperature  and  the  abundance  of  food,  and  it  is  re- 
tarded by  weather  and  scarcity  of  nourishment.  The  foregoing 
factors  are  not,  however,  the  only  causes  of  slow  development. 
I  noted  in  my  experiments  that  the  growth  of  specimens  varies 


376  The  University  Science  Bulletin. 

under  identical  external  conditions.  Very  often  a  number  of  the 
specimens  attain  full  size,  metamorphose,  and  produce  young  long 
before  others  of  the  same  generation.  The  majority,  however,  ma- 
ture in  about  five  months. 

THE   EGG. 

The  eggs  are  about  %  mm.  in  length  and  about  Yq  mm.  in  width. 
They  are  oblong  in  shape  and  slightly  arched.  Not  uncommonly 
they  were  found  adhered  together  in  pairs.  One  end  of  the  egg 
usually  has  hair  or  threadlike  projections.  Each  egg  has  a  number 
of  ridges  running  lengthwise  and  crosswise  (see  plate  XL).  The 
eggs  are  whitish  in  color,  translucent,  and  the  surface  appears  rough, 
and  is  of  delicate  skin  covering  the  aqueous  interior.  It  is  easily 
broken.  The  filaments  described  at  the  end  of  the  egg  adhere  to 
any  object  with  which  they  come  in  contact. 

MOLTING. 

I  found  without  exception  that  there  is  much  variation  in  the 
rate  of  molting  and  the  number  of  larval  skins  shed  by  the  different 
individuals  of  this  species.  Under  normal  conditions  the  larvae  molt 
twice  in  about  every  two  weeks.  Many  peculiarities  are  worthy  of 
mention.  The  same  specimen  often  sheds  its  skin  irregularly,  some- 
times within  ten  days,  and  again,  under  the  same  conditions,  not 
until  a  period  of  three  or  four  weeks  has  lapsed.  The  rapidly  grow- 
ing individuals  molt  more  frequently  than  do  those  which  have 
about  attained  their  full  size.  The  specimens  that  are  slow  in  de- 
velopment molt  less  frequently  than  do  the  larvae  which  develop 
at  the  average  rate.  The  full-grown  larvae  previously  spoken  of, 
which  continue  to  live  for  a  long  time  before  entering  the  pupal 
stage,  have  a  decidedly  slow  rate  of  molting.  The  average  rate  is 
about  once  in  every  three  weeks,  and  there  is  a  gradual  decrease 
as  the  specimen  grows  older. 

Thus  I  have  found  in  all  my  observations  that  the  number  of 
molts  is  by  no  means  constant.  As  previously  mentioned,  the  ma- 
jority of  specimens  which  complete  their  life  history  in  about  five 
months,  shed  their  skins  from  ten  to  fifteen  times,  whereas  many 
of  the  individuals  with  the  prolonged  larval  history  molt  as  often 
as  twenty  times. 

The  larva?  never  eat  their  own  skins,  nor  the  skins  of  other  indi- 
viduals of  this  species,  even  though  they  may  be  in  the  most  extreme 
stage  of  starvation.  This  was  conclusively  proven  by  placing  speci- 
mens in  a  glass  vial  for  the  purpose  of  starving  the  larvae,  and  after 


Beyer:   Trogoderma  inclusa.  377 

several  months  of  starvation,  during  which  the  larvae  had  molted 
several  times,  the  skins  were  never  attacked.  This  fact  was  also 
proven  in  the  collar  dissections  which  I  made.  In  one  collar  in 
particular,  which  had  been  kept  in  stock  for  three  years  after  the 
infestation  was  noticed,  nearly  all'  of  the  rye  seed  had  been  de- 
voured, and  the  straw  and  inner  surface  of  the  collar  were  literally 
full  of  molted  skins,  and  a  large  percentage  of  the  larvae  had  escaped 
from  the  collar. 

Just  before  molting  the  specimens  become  inactive,  and  a  break 
appears  in  the  larval  skin  along  the  median  dorsal  line.  This  ex- 
tends from  the  head  along  the  thorax  and  partly  down  the  abdo- 
men (see  plate  XL).  The  larva  assumes  a  semicircular  position, 
which  permits  the  extrication  of  the  thorax  and  the  head.  The  legs 
are  then  pulled  out  of  their  covering,  and  the  light-colored  larva 
crawls  out  of  the  exuvia.  Its  soft  covering  soon  hardens  and  be- 
comes chitinous,  and  within  a  few  hours  assumes  the  natural  yel- 
lowish-brown color. 

PUPATION. 

When  the  larva  reaches  full  growth  the  pupa  begins  to  form 
within  the  last  larval  skin.  This  is  noticeable  by  the  size,  shape  and 
the  lack  of  movement  or  locomotion.  The  pupa  is  slightly  shorter, 
larger  in  diameter,  and  apparently  makes  no  movements.  Four  or 
five  days  later  the  skin  splits  down  the  median  dorsal  line  and  the 
light-yellowish  pupa  is  exposed.  The  period  of  molting  lasts  from 
ten  to  twenty  days  at  ordinary  room  temperature.  When  the  insects 
are  fully  developed  they  emerge  through  the  large  dorsal  opening 
of  the  pupal  skin.  Should  a  specimen  be  forced  out  of  the  larval 
case  when  not  fully  matured,  though  capable  of  locomotion,  it  in- 
variably returns  to  its  former  position  within  the  protective  larval 
skin  upon  coming  in  contact  with  it.  The  pupae  upon  emerging 
are  of  a  whitish  color ;  then  comes  the  darkening  of  the  hair  on  the 
thorax  and  elytra.  In  three  or  four  days  the  thorax  and  elytra  take 
on  a  reddish  color,  commencing  at  the  thorax  and  shading  back. 
The  female  remains  in  the  pupa  case  a  day  or  two  longer  than  the 
male.    The  average  life  of  the  adult  is  about  eighteen  days. 

FOOD    HABITS. 

I  have  found  that  this  species  can  subsist  upon  a  large  variety  of 
substances.  In  considering  the  relative  value  of  some  of  the  sub- 
stances as  food  for  the  larva,  I  found  that  the  pest  apparently 
thrives  best  on  cereals.  As  was  formerly  stated,  the  larvae  were,  in 
their  natural  state,  in  almost  every  instance  found  living  on  the 


378  The  University  Science  Bulletin. 

grain  in  the  horse  collars.  Where  the  grain  was  most  abundant  1 
found  the  largest  percentage  of  larvae.  In  wandering  in  search  of 
food  many  holes  were  made  through  the  straw,  upon  which  they  fed 
in  the  meantime ;  and  upon  coming  in  contact  with  the  inner  surface 
of  the  leather,  holes  about  the  size  of  a  pinhead  were  made,  through 
which  they  emerged,  thus  injuring  the  salability  and  market  value 
of  the  leather  goods.  I  placed  a  number  of  the  specimens,  soon 
after  they  were  hatched,  on  a  leather  diet,  and  I  found  they  did  not 
eat  at  all. 

Mr.  J.  E.  Wodsedalek  (1912)  says  with  reference  to  the  species 
Trogoderma  tarsale: 

"A  number  of  the  specimens  were  placed  on  a  feather  diet,  and  although 
they  are  now  two  years  old,  they  have  grown  but  veiy  little.  When  they  were 
one  year  old  they  were  veiy  little  larger  than  the  newly  hatched  individuals, 
and  at  the  end  of  the  second  year  of  life  they  reached  a  meager  size  equal  to 
that  which  specimens  fed  on  insects  ordinarily  attain  in  two  weeks.  Their  de- 
velopment on  wool  is  even  slower." 

F.  H.  Chittenden  (1897)  states: 

"One  jar  of  flaxseed  from  the  museum  department  is  infested  chiefly  by  this 
common  museum  pest.  Many  of  the  larvae  may  be  seen  through  the  glass,  and 
large  patches  of  their  yellowish-brown  gnawings  and  excrement  show  whene 
they  have  been  at  work.    In  castor  beans  a  few  were  present. 

"That  this  species  of  Trogoderma  can  subsist  on  a  vegetable  diet  is  as  posi- 
tive as  it  is  surprising.  No  other  Coleoptera,  to  my  knowledge,  live  on  oil 
seeds,  and  I  had  nearly  arrived  at  the  conclusion  that  this  form  of  matter  was 
the  nearest  approach  to  animal  food  available,  and  that  these  insects  could 
only  thrive  on  such  vegetable  substances  as  contain  a  considerable  portion  of 
oleaginous  matter.  Judge  my  astonishment  when  a  few  weeks  after  the  dis- 
covery of  the  Trogoderma  living  in  oil  seeds,  Doctor  Howard  brought  me  a 
box  nearly  full  of  cayenne  pepper  in  which  were  several  Trogoderma  larvae. 
The  most  careful  search  failed  to  show  even  a  fragment  of  that  well-known 
red  pepper  pest,  Sitodrepa  panicea,  or  of  any  other  insect  than  the  Derme- 
stida?.  Subsequently  the  adult  was  reared  and  proved  to  be  Trogoderma  tar- 
sale. 

"To  ascertain  whether  this  species  would  breed  on  so  pungent  a  substance 
as  cayenne  pepper,  a  few  adults  were  confined  with  a  quantity  of  this  condi- 
ment. In  due  time  larvae  appeared,  and  when  examined,  August  20,  or  nearly 
ten  weeks  from  the  time  the  eggs  were  deposited,  were  in  vigorous  condition, 
the  average  individual  measuring  a  tenth  of  an  inch  in  length,  or  about  half 
that  of  the  full-grown  larvae.  Toward  the  end  of  September,  while  passing 
through  the  museum  of  this  department,  my  attention  was  attracted  by  an 
accumulation  of  powder  and  dust  about  the  edges  of  an  exhibit  of  peanut  oil 
cake,  and  another  of  Indian-turnip  bulbs.  A  large  number  of  the  larvae  and 
their  cast  skins  were  found  under  the  cakes,  also  in  the  flour  and  meal  prepared 
from  peanuts.  The  Indian-turnip  bulbs  were  very  old  and  dry,  and  might  have 
been  on  exhibition  twenty  years  or  more. 


Beyer:    Trogoderma  inclusa.  379 

"When  this  irijioct  infests  a  siibstance  of  similar  color  and  consistency  to 
Hour  and  meal,  only  a  few  laiv;^.  are  sufficient,  on  account  of  their  extraor- 
(.linary  habit  of  frequently  molting,  to  occasion  alarm.  In  fact,  appearances 
are  much  worse  than  the  reality.  Thus  in  a  small  jar  of  peanut  meal  in  which 
these  larv*  had  taken  up  their  abode,  about  forty  larval  skins  had  accumu- 
lated when  examined  Sejitembcr  27.  completely  covering  one-half  of  the  sur- 
face of  the  meal,  and  giving  the  impression  of  a  whole  colony  of  insects. 

"While  the  division  of  entomology  was  moving  into  new  quartere  a  bag  of 
Saskatchewan  spring  wheat,  formerly  kept  in  stock  for  distribution,  and  de- 
scribed on  the  label  as  a  hard,  amber  variety  with  an  exceedingly  heavy  grain, 
was  unearthed,  in  which  the  lan'se  of  this  insect  was  livdng,  three  being  present 
and  no  other  insects  except  a  colon\-  of  Arithrenus  and  a  single  stray  Silvanus. 
In  fact,  this  grain  is  so  hard  and  flinty  that  weevils  would  not  flourish  on  it. 
Soon  afterwards  I  found  larva;  in  another  lot  of  wheat  infested  with  Silvanus 
and  in  corn  containing  Calandra  crryza  and  other  small  beetles.  About  the 
same  time,  Mr.  Frank  Benton  brought  me  some  larvae  found  in  beehives, 
where  they  apparently  fed  upon  propolis  in  bee  glue.  There  are  sexeral  re- 
corded instances  of  Dermesies  lardarius  feeding  upon  wax,  or,  more  properly 
speaking,  honeycombs,  and  it  is  therefore  fairly  certain  that  Trogoderma  has 
the  same  habit,  although  not  previously  reported  in  beeliives. 

"Among  the  divi.sional  notes  I  find  one  recording  the  receipt  of  six  larvae 
of  this  species  in  a  box  of  red  pepper  from  a  correspondent  in  Utah,  Novem- 
ber 22,  1882.  These  lar\a?  were  kept  in  a  box  of  red  pepper  for  a  year,  at 
which  time  fifty-four  cast  skins  were  noticed.  The  box  was  examined  January 
14,  1887,  or  over  four  yeare  from  the  time  of  its  receipt,  when  two  larvae  and 
seventy  more  cast  skins  were  found,  but  no  trace  of  beetles,  although  it  had 
been  kept  closed  so  that  it  was  impossible  for  either  laiTse  or  adidts  to  escape. 
It  is  very  ob^■ious  that  four  larvae,  or  the  beetles  that  develoi)ed  from  them, 
had  died  in  the  interim  and  were  devoured  by  their  fellows.  In  any  case,  the 
achdt  was  not  reared,  and  no  published  statement  was  made  of  the  lar\-8e  hav- 
ing been  foimd  living  in  the  condiment. 

"The  capability  of  this  species  breeding  in  other  seeds  was  demonstrated 
by  the  discovery  of  the  larvae  living  upon  'kolu,'  an  edible  leguminous  seed 
somewhat  resembling  a  cowpea.  The  insect  had  evidently  been  first  attracted 
by  the  dead  bodies  of  the  original  inhabitants  of  the  seeds,  the  weevil,  Bruchus 
chinensis,  but  had  afterward  fed  upon  the  seeds,  even  hollowing  them  out  and 
leaving  only  the  empty  shells.  In  a  similar  manner,  larvae  were  found,  to- 
gether with  those  of  AttayenuH,  in  millet  and  pumpkin  seeds  that  had  formerly 
been  inhabited  by  the  polyphagous  Indian  meal  moth.  Plodia  interpunctella. 

"In  the  case  of  the  six  lan-ae  found  in  the  red  pepper,  it  is  not  likely  that 
four  of  them  metamorphosed,  because  if  they  had  it  is  certain  they  would 
have  been  devoured  by  their  fellows.  The  hard,  chitinous  covering  and  the 
elytra  are  never  completely  devoured,  even  by  star\'ing  specimens.  It  is  much 
more  pi-obable  that  they  died  in  the  larval  stage,  and  were  later  de\oured  by 
the  other  two  larvae,  or  they  might  have  shriveled  up  and  darkened,  and  were 
thus  easily  overlooked.  That  the  two  larvae  which  were  present  four  years 
later  were  two  of  the  original  six  is  highly  probable.  There  are  several  larvae 
in  our  laboratory  which  were  obtained  three  years  ago.  when  they  were  full 
grown,  and  they  have  not  changed  any  since." 


380  The  University  Science  Bulletin. 

behavior.  . 

Naturally  the  larvae  manifest  a  strong  negative  reaction  to  light, 
and  make  effort  immediately  after  hatching  and  when  disturbed  to 
seek  a  shaded  or  other  place  of  concealment.  If  placed  near  a  light 
or  window  they  soon  begin  to  crawl  away  from  the  light.  It  is 
also  quite  noticable  when  the  specimens  are  placed  in  a  dark  room 
and  a  strong  light  is  introduced  at  one  end  of  the  glass  dish  con- 
tainer. This  negative  phototactic  reaction  persists  throughout  life. 
It  is  at  its  highest  sensibility  to  light  just  before  pupation.  Thus 
pupse  are  most  frequently  found  in  shaded  or  dark  places  which 
afford  them  a  favorable  means  of  protection.  The  adults,  both  male 
and  female,  retain  their  negative  reaction  to  light  after  emerging 
from  their  pupal  skins.  During  the  period  of  sexual  excitment  which 
follows  a  day  or  two  later,  the  insects  are  still  negative,  and  the 
females  remain  decidedly  so  until  their  eggs  are  safely  deposited. 
Several  hours  or  a  day  after  egg-laying,  they  gradually  become 
indifferent  to  light,  and  finally  a  complete  reversal  of  their  former 
reaction  follows.  The  males  also  become  positively  phototactic 
during  the  last  days  of  their  lives.  The  larvae  of  all  stages  feign 
death  upon  being  disturbed.  However,  when  disturbance  is  con- 
tinued from  a  few  seconds  to  a  minute  at  the  most,  they  no  longer 
respond  in  the  same  manner.  When  disturbed  the  adult  insects 
make  themselves  very  compact  by  drawing  thorax  up  close  to  the 
rest  of  body.  The  head  is  drawn  upward  and  under  the  thorax,  legs 
and  antennae  are  folded  up,  and  death  is  feigned  a  considerably 
longer  time  than  in  the  larval  state.  The  average  feint  lasts  from 
one  to  ten  minutes 

VARIATION   IN    SIZE. 

The  adult  male  insects  are  smaller,  as  a  general  rule,  than  the 
female  insects,  but  the  small  individuals  are  not  always  males. 
There  is  much  variation  in  the  size  of  the  adults.  They  are  from 
1.5  mm.  to  4  mm.  in  length,  the  width  also  being  proportionate.  It 
is  difficult  to  determine  the  exact  cause  for  this  variation  in  size. 
Poor  nutrition  evidently  has  effect  upon  the  size.  However,  small 
individuals  appear  among  the  large  ones  which  have  lived  under 
very  favorable  conditions.  I  noted  marked  variations  in  the  size  of 
the  different  larvae  of  the  same  brood  within  a  day  or  two  after 
hatching.  I  observed  the  fact,  however,  that  the  small,  slowly  de- 
veloping larvae  do  not  always  produce  small  adults. 


Beyer:   Trogoderma  inclusa.  381 

adaptation  to  food  supply. 

An  interesting  phase  of  the  study  of  the  life  history  was  the  ex- 
tremely long  period  of  time  the  larvae  can  sustain  themselves  with- 
out food.  I  placed  forty  larvae  in  Petri  dishes^  ten  representative 
stages,  varying  from  newly  hatched  to  full-grown  individuals,  with- 
out any  food  whatsoever,  for  the  purpose  of  determining  the  period 
required  to  produce  starvation.  I  also  added  a  number  of  Petri 
dishes,  each  containing  one  larva  ranging  from  1  to  6  mm.  in  length, 
and  another  Petri  dish  was  added,  containing  a  number  of  a  definite 
size,  to  determine  whether  they  would  eat  their  skins  or  not.  Meas- 
urements were  made  of  all  the  individuals  and  records  kept.  Dishes 
were  examined  regularly  and  measurements  made  of  the  representa- 
tive stages.  I  also  made  a  record  of  the  cast  skins.  I  found  that 
the  larvae  never  devour  the  molted  skins  of  themselves  or  other  speci- 
mens. I  detected  no  evidence  of  cannibalism  among  the  larvaB,  even 
the  full-grown  starving  specimens  never  attacking  the  much  smaller 
individuals.  Practically  all  of  the  larvae  shed  their  skins  shortly 
after  being  deprived  of  food,  but  the  molting  process  from  this  on 
was  very  much  slower.  The  measurements  showed  in  all  cases  that 
the  different  larvae  decreased  in  size  about  one-half  their  normal 
length  after  eight  months  of  life  at  a  temperature  of  70  degrees  and 
in  ordinary  daylight.  The  newly  hatched  began  to  die  when  about 
three  months  old.  The  larvae  of  the  middle  stage  up  to  the  adult 
stage  were  all  still  surviving,  and  judging  from  existing  circum- 
stances, the  survival,  especially  of  the  full-grown  larvae,  would  be 
considerably  over  one  year.  Experiments  were  carried  on  with 
reference  to  different  kinds  of  diets  or  foods  taken  from  the  collar, 
as  rye  seed,  rye  straw  and  leather,  and  in  drawing  comparisons  it  was 
found  that  they  thrived  and  grew  rapidly  upon  the  rye  seed,  but 
refused  any  of  the  other  materials  contained  in  the  make-up  of  the 
collar. 

CONTROL  MEASURES. 
HEAT. 

I  first  took  the  trouble  to  look  over  the  field  of  available  literature 
relative  to  the  control  of  this  class  of  insects.  I  found  that  the 
French  were  the  first  to  know  the  value  of  heat,  and  to  devise  con- 
trivances for  the  heating  of  infested  buildings.  Experiments  were 
made  by  Professor  Webster  to  ascertain  the  amount  of  heat  required 
to  destroy  the  Angoumois  grain  moth,  which  gave  the  following 
results : 


382  The  University  Science  Bulletin. 

"A  temperature  of  140  degrees  continued  for  nine  hours  literally  cooks  the 
larvse  or  pupse,  a  temperature  of  130  degrees  for  five  hours  is  fatal,  as  is  also  120 
degrees  for  four  hours,  while  110  for  six  hours  was  only  partially  effective." 

It  was  also  found  that  wheat  could  be  subjected  to  a  temperature 
of  150  degrees  for  eight  hours  without  impairing  its  germinating 
properties.  In  the  second  report  of  the  state  entomologist  of  New 
York,  Prof.  J.  A.  Lintner,  speaking  of  Tribolium  jerrugineum  infest- 
ing grain  and  flour,  says : 

"A  naoderate  degree  of  heat,  120  to  130  degrees,  continued  for  a  few  hours, 
would  in  all  piobability  suffice  to  kill  all  the  eggs,  larvse  and  pupse  in  the 
material,  while  a  higher  temperature,  perhaps  150  degrees  or  more,  would  be 
needed  for  the  beetles." 

Professor  Chittenden,  in  his  paper  on  "Insects  Injurious  to  Stored 
Grain,"  states: 

'"Prior  to  the  adoption  of  carbon  disulphide  as  a  fumigant,  heat  was  relied 
upon  in  the  destruction  of  these  insects.  A  temperature  of  from  125  degrees 
to  140  degrees  Fahrenheit  continued  for  a  few  hours  is  fatal  to  grain  insects, 
and  wheat  can  be  subjected  to  a  temperature  of  150  for  a  short  time  without 
destroying  its  germinating  power." 

A  large  number  of  the  experiments  of  this  nature  were  made  rela- 
tive to  the  discovery  of  a  method  to  destroy  grain  moth,  and  from 
the  results  of  these  experiments  many  of  the  grain  insects  could 
probably  be  destroyed  in  the  same  manner,  but  it  would  require  a 
higher  temperature  to  destroy  the  adults  than  the  larvae  or  pupse. 

In  the  first  experiment  about  thirty  of  the  adults  and  larvse  were 
placed  in  a  Petri  dish  which  contained  rye  seed.  A  thermometer 
was  placed  in  the  vial,  with  the  bulb  resting  in  the  middle  of  the 
rye,  in  the  Petri  dish  containing  the  rye  grain  and  various  stages  of 
Trogoderma  inclusa.  The  Petri  dish  was  then  placed  in  a  dry-heat 
oven.  The  bottom  and  surface  of  the  interior  of  the  oven  was 
covered  with  asbestos,  and  the  Petri  dish  was  placed  upon  it,  to 
allow  uniform  heating.  The  heat  of  the  oven  was  raised  to  86  de- 
grees Fahrenheit  before  proceeding  with  the  experiment.  No  change 
was  noted  in  the  action  of  the  insects.  At  a  temperature  of  100  de- 
grees the  adults  and  larvae  began  to  crawl  out  of  the  grain;  at  a 
temperature  of  110  degrees  both  larvse  and  adults  manifested  ex- 
cessive excitement,  and  were  making  every  effort  to  escape.  They 
continued  to  be  cjuite  active  until  the  temperature  of  115  degrees 
was  reached.  At  this  temperature  the  adults  and  larvse  became  less 
active,  and  at  a  temperature  of  118  degrees  the  adults  were  all  dead, 
and  also  a  large  number  of  the  larvse.  At  a  temperature  of  119 
degrees  there  was  no  sign  of  life.    To  be  sure  that  I  had  killed  all  of 


Beykk:    Trogoderma  inclisa.  383 

the  insects,  I  raised  the  temperature  to  120  degrees,  and  then  the 
insects  were  removed  and  pUiced  in  an  incubator  and  given  a  chance 
to  recover,  but  the  test  showed  finally  that  they  were  all  dead.  I 
repeated  the  experiment  a  number  of  times,  and  found  that  as  soon 
as  a  temperature  of  from  119  to  120  degrees  was  recorded  it  proved 
fatal  to  all  stages  of  the  insect.  It  required  about  thirty  minutes 
to  reach  this  temperature. 

In  a  second  series  of  experiments  I  continued  to  use  the  dry-heat 
oven,  again  raising  the  temperature  to  86  degrees.  I  then  took  one 
of  the  infested  horse  collars  and  made  a  hole  in  the  leather,  through 
whicli  I  inserted  the  bulb  of  a  thermometer  into  the  interior  of  the 
stuffing.  The  collar  was  then  placed  in  the  oven  on  an  asbestos  floor 
to  allow  equal  heating.  The  temperature  was  gradually  raised  to 
120  degrees,  which  took  about  ten  hours.  After  leaving  the  collar 
in  the  oven  for  a  period  of  ten  hours  it  was  removed  and  placed  in 
the  incubator  to  allow  a  chance  for  the  insects  to  develop  again.  A 
day  or  two  following  I  removed  the  collar  and  proceeded  to  make 
a  thorough  dissection  of  it.  I  found  that  the  larvae  and  adults  were 
all  dead.  Upon  further  incubation  of  the  straw  stuffing  it  was  found 
that  the  eggs  had  also  been  destroyed  by  the  maximum  temperature, 
as  no  more  of  the  insects  were  hatched. 

After  demonstrating  in  the  laboratory  by  experiments  that  this 
species  of  insect  could  be  destroyed  at  a  temperature  below  that 
which  would  be  injurious  to  the  leather  of  the  horse  collars,  steps 
were  taken  to  test  out  its  results  as  to  the  practibility  in  a  seriously 
infested  collar  factory  overrun  with  Trogaderma  inclusa,  of  the 
results  which  were  demonstrated  in  the  laboratory. 

The  means  of  extermination  was  left  in  the  hands  of  the  experi- 
menter. The  collar  warehouse  w^as  the  only  infested  floor,  and  it 
was  located  on  the  sixth  floor  of  the  factory.  The  heating  facilities 
were  too  inadequate  to  produce  the  temperature  required  to  ex- 
terminate the  infestation;  hence,  according  to  instruction,  a  small 
room,  twelve  feet  long,  eight  wade  and  twelve  high,  was  constructed, 
and  lined  on  the  interior  with  asbestos.  Two  large  steam-heat  radi- 
ators were  installed,  and  the  collars  hung  on  brackets  about  the 
room.  One  large  thermometer  was  placed  in  the  room  and  several 
smaller  ones  inserted  in  the  collars.  The  results  of  this  work  proved 
successful. 

In  this  experiment  also  two  thermometers  were  employed.  One 
was  placed  in  a  collar  and  the  other  in  the  chamber.  By  recording 
the  temperatures  at  short  intervals  and  plotting  them  on  a  chart, 


384  The  University  Science  Bulletin. 

it  was  evident  that  the  temperature  rise  in  the  collar  lagged  behind 
that  of  the  chamber.  This  lag  increased  as  the  temperature  rose 
until  at  122°  F  'f  here  was  a  difference  of  nearly  two  hours.  Thus 
the  steam  was  turned  into  the  radiators  at  8  a.  m.  and  while  the 
chamber  temperature  reached  122°  F.  at  about  4  p.  m.  the  collar 
temperature  did  not  reach  this  until  6  p.  m. 

COLD. 

A  temperature  control  machine  such  as  is  used  by  this  depart- 
ment was  used  to  demonstrate  the  possibilities  of  a  freezing  tem- 
perature as  a  factor  in  the  means  of  exterminating  the  pest.  A 
number  of  the  specimens  were  placed  in  a  vial  in  the  temperature 
machine.  The  temperature  was  reduced  to  two  degrees  below  zero 
and  held  constant  for  ten  hours.  The  larvae  were  apparently  dead, 
but  after  a  short  period  of  incubation  the  specimens  began  to  be 
active  again.  I  did  not  continue  the  experiments,  as  I  realized  the 
impracticability  of  this  means  of  extermination. 

CARBON    DISULPHIDE. 

Carbon  disulphide  is  a  foul-smelling  liquid  that  volatilizes  readily 
at  ordinary  temperature,  and  produces  a  heavy  vapor  that  is 
deadly  to  insects  of  all  kinds  when  they  are  confined  in  a  closed 
space  and  must  breathe  it.  In  reading  the  data  of  the  various  books 
and  bulletins  at  hand  on  fumigation,  it  was  found  that  this  gas 
is  especially  useful  against  species  infesting  stored  grains  and  mills, 
etc.,  overrun  with  Dermestidse,  moth,  and  so  on.  There  was  reason 
to  believe,  then,  that  the  insects  within  the  collars  could  be  ex- 
terminated if  the  carbon  disulphide  could  be  properly  administered 
to  the  interior  of  the  collar. 

I  made  a  carbon  disulphide  extermination  test  of  the  Trogoderma 
inclusa  as  they  occurred  in  the  horse  collars.  I  prepared  a  box 
for  inclosing  one  of  the  collars  by  covering  it  with  paper  on  the 
inside,  and  closing  it  with  a  tight  cover.  A  small  amount  of  carbon 
disulphide  was  injected  into  the  collar  by  means  of  a  specially 
devised  syringe.  The  syringe  used  is  modeled  after  the  ordinary 
type  with  the  exception  that  a  nozzle  about  four  inches  in  length 
was  devised  to  reach  all  parts  of  the  interior  of  the  collar.  The  in- 
strument is  made  of  steel  for  durability  and  strength,  as  consider- 
able force  is  required  in  making  the  insertion.  The  nozzle  of  the 
syringe  was  inserted  in  the  collar  stuffing,  entrance  being  made  be- 
tween the  seams;  thus  there  was  no  injury  to  the  collar.  It  was 
injected  at  spaces  of  about  four  or  five  inches  apart.    From  six  to 


Beyer:   Trogoderma  inclusa.  385 

eight  drams  were  injected  into  each  collar.  The  collar  was  then 
inclosed  in  the  box  and  left  for  twenty-four  hours.  After  making 
a  thorough  dissection  of  the  collar,  the  various  larval  stages  found 
were  all  dead,  and  upon  incubation  no  evidence  of  recovery  was 
noted  in  any  of  the  larva?  or  adults. 

HYDROCYANIC-ACID   GAS. 

Hydrocyanic-acid  gas  is  a  vapor  very  destructive  to  all  life.  The 
gas  is  produced  by  adding  potassium  cyanide  to  sulphuric  acid.  I 
used  these  in  the  following  proportions: 

Potassium  cyanide,  98  per  cent  pure 1  oz. 

Sulphuric  acid,  specific  gravity  1.83 2  oz. 

Water   4  oz. 

I  placed  several  of  the  infested  collars  in  a  closed  chemistry  hood. 
I  put  an  earthen  vessel  inside,  containing  water,  poured  the  sul- 
phuric acid  slowly  in  the  water,  and  then  added  the  potassium  cya- 
nide, and  immediately  closed  the  hood  and  left  the  collars  exposed 
to  the  hydrocyanic  acid  for  two  hours,  and  then  opened  the  hood 
and  let  the  gas  escape.  I  then  took  the  collars  from  under  th-e 
hood  and  examined  the  stuffing.  There  was  no  trace  of  life  or  re- 
covery of  the  insects  in  the  different  stages. 

SUMMARY  AND  CONCLUSION. 

The  Trogoderma  inclitsa  discussed  in  this  paper  were  found  in 
horse  collars  that  were  sent  to  this  department,  requesting  our  ad- 
vice and  assistance  in  determining  the  kind  and  source  of  infesta- 
tion and  measures  to  be  used  in  the  control  of  this  damaging  insect. 

With  regard  to  the  source  of  infestation:  Upon  opening  the  col- 
lars and  finding  grain  in  the  straw  stuffing,  and  upon  examination 
of  the  grain  in  many  instances,  I  noted  that  it  had  been  eaten  upon 
by  the  larvae  of  this  insect,  and  through  the  instrumentality  of  my 
series  of  investigations  in  relation  to  the  life  history,  I  found  that 
they  thrived  much  better  upon  the  grain  than  any  other  material 
found  in  the  make-up  of  the  collars.  This  gave  me  a  clue  as  to  the 
source  of  infestation.  After  getting  some  of  the  stored  unthreshed 
rye,  and  some  of  the  rye  seed  which  had  been  stored  where  it  had 
been  raised,  I  found  that  a  series  of  incubations  produced  exactly 
the  same  species  that  was  found  in  the  collars. 

In  considering  the  matter  of  infestation  and  the  measures  to  be 
used  for  its  control,  I  suggest  that  the  grain  be  eliminated  as  nearly 
as  possible  from  the  straw  to  be  used  in  the  stuffing  of  the  collars, 
and  there  will  be  no  liability  of  serious  infestation.    However,  as  I 


386  The  University  Science  Bulletin. 

have  experienced  personally  and  through  correspondence;  many  of 
the  people  engaged  in  the  manufacture  of  horse  collars  are  not  aware 
of  the  value  of  clean-threshed  straw,  the  seed  of  which  forms  an 
abundant  food  supply  for  these  insects,  and  thus  follows  the  dilemma 
caused  by  the  infestation.  Applicable  to  such  cases,  I  have  per- 
formed a  series  of  experiments  to  demonstrate  their  relative  value 
concerning  the  means  of  extermination.  Experiments  were  carried 
on  with  heat  and  cold  temperatures  and  carbon  disulphide  and 
hydrocyanic-acid  gases,  and  in  conclusion  I  feel  at  liberty  to  state 
that  heat  is  to  be  preferred  as  the  best  means  of  control.  It  insures 
extermination,  and  is  by  far  the  most  economical  as  well  as  the 
safest  means  of  eradicating  the  infestations. 

With  reference  to  the  matter  of  multiplicity  and  distribution  of 
this  species  of  insect,  it  was  found  to  be  a  rare  species  generally 
distributed  over  the  the  United  States  and  Europe,  and  is  omniv- 
orous in  its  feeding  habits.  I  found  this  species  to  thrive  and  de- 
velop much  more  rapidly  upon  seeds  and  grain  foods  than  any  other 
available  material  which  I  had  at  hand  for  testing  out  the  food 
habits. 

In  making  a  summary  of  the  variations  of  the  life  history  of  a 
number  of  individuals  of  the  same  generation,  I  noted: 

1.  The  adults  oviposit  from  four  to  six  days  after  emergence. 

2.  The  number  of  eggs  laid  by  different  individuals  varied  from 
ten  to  forty-five. 

3.  The  eggs  hatch  in  from  eiglit  to  twelve  days  at  ordinary  room 
temperature. 

4.  The  larval  life  lasts  about  five  months,  on  the  average. 

5.  The  time  of  pupation  is  from  ten  to  fourteen  days. 

6.  The  adult  lives  from  eight  to  twenty-five  days. 

BIBLIOGRAPHY. 

F.  H.  Chittenden.     1893. — Herbiverous  Habits  of  Certain  Dermestids.    Bull. 

2,  N.  S.  Div.  Ent.,  U.  S.  Dept.  Agr.,  i)p.  36.  37.   - 
1897. — Granivorous  and  Other  Habits  of  Certain  Dermestids.    Bull.  8  N.  S. 

Div.  Ent.  U.  S.,  pp.  14-24.  fig.  1. 
L.  O.  Howard.    Extract  from  Corres.  Bull.  44,  Div.  Ent,  U.  S.  Dept.  Agr.,  Apr., 

pp.  90-99. 
H.  F.  Jayne.    1882.    Revision  of  the  Derm,  of  the  U.  S.    Proc.  Amer.  Philos. 

Soc,  vol.  XX. 
C.  V.  Riley.     1883.—Trugoderma  as  a  Museum  Pest.     Amer.  Nat.,  vol.  17 

p.  199. 
1S83. — Number  of  Molts  and  Length  of  Lar\-al  liife  as  Influenced  by  Food. 

Amer.  Nat.,  vol.  17,  pp.  347-548. 


Beyer:    Trogoderma  inclusa.  387 

F.  H.  Snow.    1SS2.—A  New  Museum  Pest.    Psyche,  vol.  3,  pp.  351,  352. 

1S94.— Insect  Life,  vol.  VI,  p.  226. 

1894- — Proceedings  of  the  Columbus  Hort.  Soc,  vol.  IX,  p.  12;  Apr. 

.?S56.— Canadian  Entomologist,  vol.  XXVIII,  p.  262;  Oct. 
W.  S.  Blatchley.    Coleoptera  of  Indiana,  p.  593. 

Dec,  1912,  Annals  of  Ento.  Soc.  of  Amer.,  p.  367. 
Le  Conte.   Synopsis  of  the  Dermestidse  of  U.  S.  Proc.  Phil.  Acad.  Nat.  Sc, 

vol.  VIII,  1854,  pp.  106-113. 
Casey.    Review  of  the  American  Dermestidse.    Jour.  N.  Y.  Ento.  Soc,  VIII, 
1900,  pp.  138-165. 


388  The  University  Science  Bulletin. 


PLATE  XXXIX. 

Photographs  of  leather  horse  collars  damaged  by  the  dermestid  Trogoderma 
inclusa  Lee.  In  figures  A  and  B,  white-headed  pins  were  inserted  to  indicate 
the  position  of  the  holes  made  by  emerging  beetles.  Figvu'e  C  shows  the  exit 
holes  made  by  the  beetles. 


PLATE  XXX IX. 


B 


(389) 


Photographs  by  P.  A.  Readio. 


PLATE  XL. 

Truyodeniia  inclusa  I^ec. 

1.  Ventral  view  of  beetle. 

2.  Dorsal  view  of  beetle. 

3.  Ventral  view  of  jnipa. 

4.  Eggs. 

5.  Antennae. 

6.  Ventral  view  of  larva. 

7.  Pupa. 

8.  Dorsal  view  of  larva. 

(390) 


Beyer:    Trogoderma  inclusa. 


391 


PLATE  XL. 


.-(rfS 


'V'hi>    8 


THE 

KANSAS  UNIVEESITY 

Science  Bulletin 


Vol.  XIV,  No.  16— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  16.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

The  Larva  of  a  Chironomid  (Diptera) P.  W.  Claassen. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  INIYERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  16. 


The  Larva  of  a  Chironomid  {Trissocladius  equitans  n. 
sp.)  Which  Is  Parasitic  upon  a  May-fly  Nymph  (Rith- 
rogena  sp.).* 

BY  p.  W.  CLAASSEN. 

IN  AUGUST,  1919,  while  spending  a  few  weeks  in  Colorado,  the 
writer  was  collecting  aquatic  insects  in  the  Big  Thompson  river 
in  Estes  Park.  This  river  is  a  typical  mountain  stream.  The  water 
is  very  cold  and  rushes  along  in  a  swift  current  over  a  stony  bed. 
Aciuatic  insects  of  the  swift-water  type  were  very  abundant.  Among 
the  May-fiy  nymphs  collected  there  were  found  about  a  dozen  speci- 
mens of  one  species  which  presented  a  curious  appearance.  Upon 
examining  some  of  these  nymphs  it  was  found  that  each  one  carried 
upon  its  back  a  large,  white  dipterous  larva.  These  larvae  had  at- 
tached themselves  to  the  thorax  of  the  nymphs  underneath  the  wing 
pads.  The  large  size  of  the  larva  forced  the  wing  pads  of  the  May- 
fly nymph  upward  at  a  very  decided  angle  and  gave  the  nymph  a 
humped-up  appearance  as  it  rested  upon  the  stone. 

All  the  collected  material  was  preserved  in  alcohol  and  taken  back 
to  Ithaca,  N.  Y.,  for  study.  When  the  material  was  examined  more 
closely  in  the  laboratory,  some  of  these  May-fly  nymphs  were  found 
to  carry  larvae,  while  others  of  the  same  species  carried  pupae  of  the 
dipterous  insect. 

The  May-fly  nymph  proved  to  be  a  species  of  the  genus  Rithro- 
gena.  Although  it  was  impossible  to  determine  definitely  the  genus 
of  the  diptcron,  Dr.  0.  A.  Johannsen,  who  examined  it,  felt  certain 
that  it  belonged  to  the  family  Chironomidse.  The  scarcity  of  mate- 
rial and  the  lack  of  adult  forms,  however,  made  it  impossible  to  sat- 

*  Read  before  the  joint  session  of  the  Entomological  Society  of  America  and  Ecological 
Society  of  America,  January,  1922.     Withdrawn   for  publication  in  this  bulletin. 

(395) 


396  The  University  Science  Bulletin. 

isfactorily  establish  the  relationship  which  existed  between  these 
two  forms. 

During  the  summer  of  1921  the  writer  made  another  trip  to  Estes 
Park,  Colo.,  and  again  found  these  May-fly  nymphs  in  the  same 
situation  in  the  Big  Thompson  river  where  they  had  previously  oc- 
curred. A  careful  search  was  then  made  over  a  distance  of  a  mile  or 
more  in  this  stream,  but  there  was  only  this  one  particular  spot  in 
which  the  Rithrogena  nymphs  could  be  found  in  considerable  num- 
bers. This  was  in  a  part  of  the  stream  where  the  current  was  quite 
swift  and  the  water  not  over  six  to  ten  inches  deep.  The  bed  of  the 
stream  was  covered  with  stones,  many  of  the  stones  projecting 
above  the  surface  of  the  water.  May-fly  nymphs,  stone-fly  nymphs, 
caddis  worms  and  other  swift-water  forms  were  very  plentiful.  As 
many  as  five  or  six  specimens  of  the  Rithrogena  nymphs  occurred  on 
a  single  stone.  Upon  taking  a  stone  out  of  the  water  and  turning  it 
over,  these  nymphs  would  quickly  glide  to  the  under  side  of  the 
stone,  appearing  to  be  much  more  elusive  than  any  of  the  other  spe- 
cies of  the  May-fly  nymphs  present.  A  total  of  nearly  300  of  the 
Rithrogena  nymphs  were  collected,  and  more  than  ninety  per  cent 
of  them  were  found  to  carry  either  a  larva  or  a  pupa  of  the  chi- 
ronomid. 

An  attempt  was  then  made  to  rear  to  the  adult  stage  the  May-fly 
and  the  chironomid.  In  order  to  accomplish  this  a  number  of  the 
nymphs  which  carried  pupsB  were  placed  in  small  wire  cylinder 
cages.  Some  of  the  cages  were  placed  in  the  part  of  the  stream 
where  the  nymphs  naturally  occurred,  while  others  were  placed  in 
a  spring  near  the  writer's  cottage,  where  they  could  be  kept  under 
close  observation.  Three  males  of  the  chironomids  emerged  on 
August  7,  two  of  them  from  cages  in  the  stream  and  one  from  a  cage 
in  the  spring.  A  female  also  emerged  on  the  same  date,  but  it  es- 
caped. On  August  8  two  males  of  the  May-fly  emerged,  one  from 
a  cage  in  the  stream  and  the  other  from  a  cage  in  the  spring.  On 
the  previous  day  two  females  of  the  May-fly  were  caught  in  a  net 
near  the  same  spot  where  all  the  material  was  collected,  and  these 
later  proved  to  belong  to  this  same  species. 

The  necessity  of  leaving  the  park  on  August  8  prevented  any  fur- 
ther rearing  work,  and  the  material  from  the  cages  was  added  to  the 
alcoholic  specimens  and  taken  back  to  the  laboratory  at  Cornell 
University. 

The  May-fly  has  been  determined  by  Dr.  J.  G.  Needham,  who 
recognizes  it  as  a  new  species  of  the  genus  Rithrogena.    A  descrip- 


Claassen:    Larva  of  a  Chironomid.  397 

tion  of  this  species  will  be  given  by  Doctor  Needham  in  a  paper 
which  is  soon  to  be  published. 

The  chironomid  is  a  new  species  of  Trissocladius,  a  genus  new  to 
this  country.* 

DESCRIPTION  OF  THE  STAGES  OF  THE  CHIRONOMID. 

Trissocladius  equitans  n.  sp. 

ADULT. 

Length,  4.5  mm.,  exclusive  of  antennae.    General  color,  blackish  brown. 

Head  blackish ;  ej-es  black,  naked,  slightly  emarginate  on  inner  margins,  the 
facets  moderately  rounded;  distance  between  the  eyes  from  above  greater 
llian  the  length  of  the  eye.  Labium  short,  thick,  and  reaching  to  the  second 
segment  of  the  palpi.  Palpi  short,  three-segmented;  the  first  segment  nearly 
twice  as  long  as  broad;  second  segment  twice  as  long  as  broad;  third  seg- 
ment graduallj'  tapering  to  the  tip  and  a  little  longer  than  second  segment; 
total  length  of  palpi  about  .25  mm.  Antennae  composed  of  fourteen  segments, 
the  terminal  one-half  again  as  long  as  the  combined  length  of  segments  two 
to  twelve;  ba^al  segment  large;  second  segment  about  twice  as  long  as  third; 
total  length  of  antenna  L3  mm.;  entire  antennae  covered  with  long  hairs,  the 
basal  hairs  as  long  as  the  terminal  segment.  Epistome  without  bristles;  trans- 
verse suture  distinct. 

Thorax  blackish  brown,  with  whitish  pruinose  patches;  dorsum  with  a 
naiTow  median  line,  depressed  in  front  and  raised  into  a  feeble  carina  behind 
the  middle;  surface  smooth  except  for  a  few  hairs  on  each  side  of  the  broad, 
flattened  longitudinal  area  of  the  mesonotum ;  collar  incised  in  middle,  angles 
rounded;  scutellum  and  metanotum  blackish,  smooth;  plura  and  pectus  black- 
ish, smooth. 

Abdomen  compressed,  blackish,  with  long,  yellowish  hairs ;  basal  segment  of 
the  male  clasper  about  twice  as  long  as  the  distal  segment,  spoon-shaped, 
the  median  projection  or  tubercle  blunt,  with  short  hairs,  basal  segment  below 
with  long  hail's;  distal  segment  narrow  at  base  and  gradually  enlarging  toward 
the  apex,  a  few  hairs  above  and  underneath  with  two  short  spines  near 
the  tip. 

Wings  milkj^  white,  reaching  to  the  base  of  the  claspers;  surface  finely 
punctate;  anal  lobe  produced  and  fringed  with  long  hairs;  venation  as  in 
figure  14. 

Halteres  pale,  slightly  infuscated. 

Legs  light  brown,  hairy;  tibia  of  front  leg  L4  times  the  length  of  metatarsus; 
a  single  distinct  spur  at  distal  end  of  tibis.  the  ones  on  the  posterior  legs 
being  much  larger;  fourth  and  fifth  tarsal  segments  of  equal  length;  tarsal 
claws  squarely  truncate,  surface  fluted;  no  pectinate  empodium  present. 

Type  in  the  Cornell  University  collection,  now  mounted  in  balsam.  Para- 
types,  two  males;  one  a  pinned  specimen,  the  other  in  alcohol,  in  the  Cornell 
University  collection. 

These  specimens  are  all  from  Estes  Park,  Colo.,  August  7,  1921. 

♦Acknowledgments  are  due  to  Dr.  O.  A.  Johannsen,  who  recognized  this  as  a  new  species, 
and  without  whose  assistance  the  following  descriptions  could  not  have  been  adequately  made. 


398  The  University  Science  Bulletin. 

LARVA. 

Length,  6-7  mm.  when  fully  grown.    Color  white. 

Head  very  small,  not  over  .25  mm.  in  diameter.  Prothoracic  proleg  double, 
short,  with  many  short  spines;  caudal  prolegs  short,  each  with  a  ring  of  about 
24  short  spines.    Anal  gills  not  apparent. 

Head  yellowish  brown,  with  a  narrow  black  hind  border;  on  the  ventral 
surface  of  the  head,  near  the  hind  border,  there  is  on  each  side  a  light  oval 
spot,  suggestive  of  an  ocellus.  Mouth  parts  very  small  and  difficult  to  dis- 
sect out;  t-hey  are  much  reduced  in  size;  the  mandibles  are  composed  of  a 
single  sharp  tooth  with  two  short  spines  on  the  inner  margin  near  the  base 
of  the  tooth;  labrum  small,  the  front  margin  excavated  in  the  middle  so  as 
to  leave  a  blunth'  rounded  chitinized  tooth  on  each  side;  labial  palpi  very 
short :  antennae  minute,  easily  overlooked. 

PUPA. 

Length,  5-5.3  mm.  Color  brown;  a  narrow,  blackish  margin  around  the 
wings,  and  a  narrow  longitudinal  dark  line  each  side  of  the  abdomen.  The 
-^ntire  pupa  is  smooth  and  devoid  of  any  vestiture.  Breathing  trumpets  not 
well  developed,  but  represented  by  a  spiracularlike  chitinized  area  each  side 
of  the  mesothorax.  These  breathing  trumpets  are  best  seen  in  the  late 
larval  stage  when  the  pupa  is  being  formed  (fig.  7).  Segment  one  to  three 
of  the  abdomen  smooth;  segments  four  and  five,  above,  each  with  a  double 
transverse  band  of  fine  spines  on  the  posterior  margin,  the  spines  of  the  hind- 
most bands  directed  forward  and  at  least  twice  as  long  as  the  spines  on  the 
band  immediately  in  front ;  segments  six,  seven  and  eight)  each  with  a  single 
transverse  band  of  short  spines.  Genital  sacs  smooth,  no  hairs  or  spines 
present.  On  the  lateral  margin  of  the  abdominal  segments  the  vestigial 
spiracles  are  visible. 

THE  BIOLOGICAL  RELATIONSHIP 

Between  Trissocladms  equitans  and  Rithrogena  sp. 

With  plentiful  material  on  hand  it  has  been  possible  to  establish 
the  relationship  which  exists  between  the  immature  stages  of  these 
insects.  The  larva  of  Trissocladius  equitans  is  parastic  upon  the 
Rithrogena  May-fly  nymph.  Although  it  has  not  been  possible  to 
learn  in  what  manner  the  larva  establishes  itself  as  a  parasite  on  the 
nymph,  indications  are  that  Trissocladius  equitans  spends  its  entire 
larval  life  as  a  parasite  upon  the  May-fly  nymph.  Different-sized 
larvae,  representing  the  instars,  except  possibly  the  first,  were  found 
under  the  wing  pads  of  the  nymphs. 

The  parasite  attaches  itself  to  the  posterior  margin  of  the  meso- 
thorax underneath  the  wing  pads,  where  it  imbeds  its  head  in  the 
softer  tissue  of  the  nymph  and  where  it  draws  its  nourishment  from 
the  host.  The  larva  spins  a  sheet  of  silk,  which  completely  invests 
its  body,  and  by  means  of  which  it  attaches  itself  firmly  to  the 
body  of  the  nymph.    At  first,  stretched  to  its  full  length,  the  larva 


Claassen:    Larva  of  a  Cpilronomid.  399 

lies  across  the  body  of  the  nymph,  but  as  it  grows  hxrger  the 
posterior  end  of  (ho  body  doubles  under,  and  later,  when  the  larva 
becomes  full  grown,  the  middle  part  of  its  body  projects  backward 
over  the  abdomen  of  the  nymph  in  the  form  of  a  letter  U.  When 
ready  to  pupate  the  larva  releases  its  hold  at  the  head  end  and  turns 
back  over  the  abdomen  of  the  nymph.  The  pupa  thus  extends  over 
about  half  the  abdomen  of  the  host.  Tlic  transparent  sheet  of  silk 
invests  the  pupa  as  well  as  the  larva.  Just  before  the  adult  is 
ready  to  emerge  the  pupa  breaks  through  this  investing  membrane 
and  rises  to  the  surface  of  the  water,  where  the  adult  emerges  in  a 
manner  similar  to  that  in  other  chironomids. 

That  the  larva  is  a  parasite  upon  the  nymph  is  apparent  from 
the  following  facts:  First,  the  silk  membrane  completely  sur- 
rounds the  larva.  There  is  no  opening  in  this  membrane  which 
would  make  it  possible  to  obtain  food  from  the  outside.  Second, 
an  examination  of  the  stomach  contents  of  a  number  of  larvae 
failed  to  reveal  the  presence  of  any  vegetable  material,  but  did  con- 
tain animal  matter,  especially  fat  bodies.  This  fact  was  also  borne 
out  in  a  study  of  cross  sections  of  the  larvse.  Third,  a  study  of 
the  head  and  mouth  parts  of  the  parasite  reveal  their  reduced  con- 
dition. Tlie  head  is  very  small  in  proportion  to  the  size  of  the  body 
and  the  mouth  parts  are  much  reduced. 

The  European  species  of  the  genus  Trissodadius,  of  which  the 
larvae  are  known,  are  all  found  as  free  living  forms  feeding  upon 
vegetable  matter.* 

The  reduced  size  of  the  head  and  mouthparts  of  Trissodadius 
equitans  indicate  that  this  parasitic  relationship  has  existed  for  a 
long  time.  Whether  the  parasite  ever  becomes  so  injurious  as  to 
kill  its  host  the  writer  has  not  been  able  to  determine.  Another 
question  of  interest  is  what  happens  to  the  parasite  when  the  nymph 
casts  its  skin.  The  life  cycle  of  the  parasite  is  either  so  short  that 
development  is  completed  during  a  nymphal  instar  of  the  host,  or 
else  the  parasite  must  be  able  to  detach  itself  from  the  cast  skin 
and  reestablish  itself  upon  the  newly  emerged  nymph  or  upon  an- 
other individual,  otherwise  it  must  perish. 

There  appears  to  be  no  relationship  between  the  relative  ages  of 
the  host  and  parasite.  Pupa?  of  the  parasite  were  found  upon 
medium-sized  nymphs,  while,  on  the  other  hand,  young  larvse  oc- 
curred upon  nymphs  that  were  evidently  in  the  last  nymphal  in- 
star. 


*Barnard,  K.  H.,  in  the  Entomologist's  Monthly  Magazine,  vol.  47:76-78,  1911,  reports 
the  case  of  a  chironomid  larva  (name  not  given)  which  is  parasitic  upon  a  fresh  water  snail, 
Limnoea  peregra. 


400  The  University  Science  Bulletin. 


PLATE  XLI. 

Fig.  1.   Rithrogena  nymph  with  a  larva  of  Trissocladius  equitans  under  the 
wing  pads. 

Fig.  2.   Nymph  with  a  pupa. 


Claassen:   Larva  of  a  Chironomid. 


401 


PLATE  XLL 


402  The  University  Science  Bulletin. 


PLATE  XLII. 

Fig.  3.   Side  view  of  May-fly  nymph,  showing  the  chironomid  pupa  under 
the  wing  pads. 

Fig.  4.   Young  larva  of  T.  equitans  under  the  wing  pads  of  the  nymph. 
Fig.  5.   Full-grown  larva  on  the  nymph. 
Fig.  6.   Pupa  on  the  nymph. 


PLATE  XIJI. 


^  fe- 


itii 

I 


A 


(403) 


PLATE  XLIII. 

Fig.  7.  Full-grown  larva  of  T.  equitans.  In  the  thorax  may  be  seen  the 
breathing  trumpets  of  the  developing  pupa  within. 

Fig.  8.    Mandible  of  the  larva  of  T.  equitans. 

Fig.  9.    Labium  of  the  larva. 

Fig.  10.    Hind  proleg  of  the  larva. 

Fig.  11.    Portion  of  tlie  abdomen  of  the  pupa. 

Fig.  12.    Adult,  T.  equitans. 

Fig.  13.    Clasper  of  male. 

Fig.  14.    Wing  of  male. 

Fig.  15.    Young  larva. 

Fig.  16.  Nearly  full-grown  larva.  The  black  line  indicates  the  edge  of  the 
silk  membrane  which  envelopes  the  lana  and  by  means  of  which  it  is 
attached  to  its  host. 

(404) 


Claassen:   Larva  of  a  Chiroxomid. 


405 


PLATE  XLIIL 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  17— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  17.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

Water  Insects  from  a  Portion  of  the  Southern  Utah 

Desert R.  C.  Moore  and  H.  B.  Hungerjord. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE,  KAN. 


Entered  at  the  post  office  in   Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  17. 


Water  Insects  from  a  Portion  of  the  Southern  Utah 

Desert. 

BY  R.  C.  MOORE, 
Professor  of  Geology,  University  of  Kansas ;   and 

H.  B.  HUNGERFORD, 
Professor  of  Entomology,  University  of  Kansas. 

INTRODUCTION. 
THE  COLORADO  PLATEAU. 

Not  the  least  interesting  of  that  well-known  and  yet  little  known 
country  of  varied  attractions,  the  Great  Western  Cordillera  of 
America,  is  the  region  of  lofty  plateaus,  towering  cliffs  and  deep, 
impassable  canyons  which  is  known  as  the  Colorado  plateau. 
Bordered  on  the  east  by  the  snow-clad  peaks  of  the  Rocky  Moun- 
tains, on  the  north  by  the  Uinta  mountains,  and  on  the  west  and 
south  by  low-lying  deserts  of  the  Great  Basin  and  the  lower  Colo- 
rado valley,  the  Colorado  plateau  includes  most  of  western  Colorado, 
eastern  and  southern  Utah,  northern  Arizona  and  northwestern 
New  Mexico.  Unlike  the  serrated  peaks,  irregular  jagged  spurs  and 
sharp-topped  divides  of  the  Rockies',  or  of  the  mountain  ranges  in 
the  Great  Basin  and  Arizona  deserts,  the  plateau  country  is  a  land 
of  elevated,  essentially  flat-topped  tables,  which  are  terminated  for 
the  most  part  in  steep,  irregularly  trending  cliffs,  and  of  great 
canyons  which,  converging  on  and  culminating  in  the  world-famous 
canyon  of  Colorado  river,  ramify  almost  every  section  of  the  plateau 
province.  The  tablelands  are  formed  by  hard  rock  formations 
which  lie  in  more  or  less  nearly  horizontal  positions,  and  the  steep 
cliffs  which  border  the  plateaus  or  wall  in  the  canyons  mark  the 
edges  of  these  hard  formations.  Exceptions  to  the  general  archi- 
tectural scheme  of  the  Colorado  plateau  country  are  a  few  small 
mountain  masses  of  igneous  origin,  volcanic  cones  like  the  San  Fran- 

(409) 


410  The  University  Science  Bulletin. 

cisco  mountains  in  Arizona,  or  laccolithic  intrusions  like  the  Henry 
mountains  in  southern  Utah.  These  are  not  important  in  the  aggre- 
gate, but  are  striking  on  account  of  contrast. 

All  of  the  Colorado  plateau  region  is  arid  or  semiarid.  There  is 
little  rain,  and  most  of  that  which  does  come  falls  during  a  small 
part  of  the  year  and  in  torrential  showers.  Much  of  the  character- 
istic topography  and  the  aspect  of  the  country  in  general  is  due 
chiefly  to  this. 

THE  HIGH  PLATEAUS  OF  SOUTHERN  UTAH. 

The  features  and  the  conditions  which  are  broadly  typical  of  the 
Colorado  plateau  as  a  whole  find  especially  characteristic  expression, 
and,  indeed,  culminate  in  the  south  central  part  of  Utah  and  adja- 
cent portion  of  Arizona.  From  the  Grand  canyon  of  Colorado  river, 
the  most  profound  and  prodigious  of  the  canyons,  the  stratified 
rock  platforms  rise  tier  on  tier  like  gigantic  stairs  ascending  north- 
ward. The  top  "stairs,"  in  southern  Utah,  have  an  elevation  of 
more  than  10,000  feet  above  sea  level,  and  comprise  the  so-called 
high  plateaus.  It  is  almost  inevitable  that  this  lofty  plateau 
country,  so  closely  adjacent  to  the  deepest  of  the  canyons,  should 
be  intricately  dissected  by  tributary  canyons.  The  high  plateaus 
and  adjoining  region  may  therefore  be  specially  designated  as  the 
"canyon  lands."  Travel  is  by  tortuous  and  extremely  toilsome 
routes,  now  in  the  depths  of  a  profound  abyss,  now  crossing  a  pla- 
teau spur  of  mountainous  proportions.  Some  areas  are  absolutely 
inaccessible,  and  large  districts  are  almost  unexplored.  Population 
is  confined  to  a  very  few  of  the  accessible  valleys  where  irrigation 
permits  cultivation  of  a  little  of  the  land  adjacent  to  the  water 
supply.  Aside  from  these  outposts  of  determined  agricultural  skill 
and  industry,  the  region  is  traversed  only  by  occasional  cattlemen, 
prospectors  or  explorers. 

CLIMATE. 

The  two  main  features  which  characterize  the  climate  of  southern 
Utah  and  affect  more  or  less  directly  the  life  of  the  region,  as  well  as 
physiographic  processes,  are  dryness  and  temperature  variation. 
In  part  of  the  area  rainfall  is  less  than  five  inches  in  the  year,  and  in 
all  of  it  the  average  is  less  than  ten  inches  per  year.  In  general, 
there  is  most  rain  in  the  three  summer  months,  July  August  and 
September;  and  least  rain  in  the  spring  months,  April,  May  and  June. 
The  fall  and  winter  months  have  an  intermediate  rainfall.  The 
effect  of  the  dry  spring  season,  when  most  plants  begin  to  develop 
rapidly  and  when  many  of  the  animals,  especially  the  insects,  pro- 


jNIoori:  and  Hiwckkford:   Water  Insects.  411 

gress  swiftly  through  early  life  stages  toward  the  vigor  of  mid- 
summer maturity,  is  to  retard  or  to  inhibit  the  development  of  life. 
Seeds  may  germinate,  but,  without  rains,  growth  lags  or  ceases. 
Larva)  of  insects,  polliwogs,  and  whatever  animal  life  is  dependent 
on  waters,  are  restricted  to  permanent  pools,  springs  or  streams,  for 
there  are  no  temporary  breeding  waters  in  the  dryness  of  the  spring. 
The  distribution  of  the  yearly  precipitation  at  four  stations  in  the 
vicinity  of  the  localities  described  in  this  paper,  is  shown  in  chart 
on  page  412. 

The  temperature  of  the  plateaus  is  influenced  to  a  large  extent 
by  the  clear,  dry  atmosphere  and  by  the  altitude.  During  the  days 
the  sun  warms  the  air  and  the  ground,  and  the  bare  rocks  reflect 
the  heat  rays  back  into  the  air.  At  night  radiation  is  rapid  and  the 
temperature  quickly  falls.  This  large  diurnal  range  in  temperature 
is  characteristic  of  arid  or  semiarid  regions.  At  Tropic,  in  central 
western  Garfield  county,  a  freezing  temperature  in  each  of  the 
twelve  months  has  been  reported,  while  the  maximum  temperature 
is  greater  than  100  degrees.     (See  chart,  page  413.) 


412 


The  University  Science  Bulletin. 


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414  The  University  Science  Bulletin.  ^    . 

surface  "waters. 

Though  the  topography  of  southern  Utah  is  very  evidently  the 
result  chiefly  of  erosion  by  running  water,  there  are  few  perennial 
streams  in  the  region.  The  Colorado,  master  stream  of  the  plateau 
country,  gathers  the  waters  from  the  west  slopes  of  the  Rockies, 
from  the  Uintas  and  other  ranges  where  rains  and  melting  snows 
furnish  a  varying  but  unending  water  supply.  The  tributaries 
which  unite  to  form  the  Colorado  begin,  for  the  most  part,  as  clear 
mountain  streams,  but  in  southern  Utah  the  river  carries  so  heavy 
a  load  of  mud  and  sand  that  the  water  is  dark  reddish  brown.  On 
this  account  the  name  Colorado  was  first  applied.  The  river  is 
large  and  swift,  with  swirling  eddies  and  numerous  rapids.  The 
only  permanent  tributaries  to  the  Colorado  in  southern  Utah  are 
San  Juan  river  on  the  south  and  Dirty  Devil  and  Escalante  rivers 
on  the  north.  Each  of  these  streams  flows  in  a  deep  canyon,  and, 
except  for  size,  is  essentially  similar  to  the  Colorado.  The  Esca- 
lante, which  is  the  main  stream  in  the  region  from  which  most  of 
the  collections  of  insects  were  obtained,  is  fed  by  the  melting  snows, 
rains  and  springs  of  the  southern  extremity  of  the  highest  of  the 
plateaus,  the  Aquarius.  Its  waters  are  somewhat  clearer  than  those 
of  the  other  streams,  because  most  of  its  course  lies  through  massive 
sandstones. 

The  numerous  smaller  streams  in  southern  Utah  contain  water 
only  at  times  of  local  rainfall.  The  run-off  is  very  rapid,  and  after 
a  torrential,  muddy  flood  of  less  than  an  hour,  or  at  most  a  few 
hours,  the  flow  ceases.  This  type  of  swift,  intermittent  streams  is 
characteristic  of  all  the  plateau  region. 

After  rains  there  are  in  places  pools  or  "tanks"  in  such  natural 
hollows  as  may  retain  any  of  the  water.  Some  of  these  are  found 
in  the  stream  channels,  others  in  eroded  depressions  in  the  nakecf 
rock.  Some  are  small,  shallow  and  short-lived  under  the  rays  of 
an  intense  sun  and  a  thirsty  atmosphere;  others  are  larger,  and 
occasionally  deep.  Those  in  the  channels  of  streams  commonly 
contain  very  muddy  water,  which,  because  of  the  excessive  fine- 
ness of  the  mud,  retains  the  sediment  in  suspension  until  evapora- 
tion gradually  gives  it  the  consistency  of  thick  soup  or  gravy,  and 
finally  of  brick.  Where  the  rain  water  accumulates  in  sandstone, 
or  where  the  gathering  waters  do  not  cross  exposures  of  the  soft 
shales  which  furnish  most  of  the  muds,  the  waters  remain  fairly 
clear.  The  larger  "tanks"  may  retain  water  from  year  to  year, 
shrinking  slowly  through   times  of   drought,  but  refilling   on  the 


Moore  AND  Hixgerford:   AVatkk  Insects.  415 

coming  of  rain.  These  little  pools  of  moderately  clear  water  afford 
sanctuary  for  such  water-breeding  or  water-living  animals  as  in- 
habit the  region,  and  in  some  cases  it  is  a  populous  and  assorted 
community  that  crowds  together. 

Springs  may  be  mentioned  among  the  surface  waters  of  southern 
Utah,  although  they  are  not  numerous,  and  the  water  from  them 
very  short h^  sinks  into  the  ground.  They  are  in  many  cases  nearly 
permanent,  and  are  an  important  source  of  supply,  especially  as 
regards  the  uses  of  man.  None  of  the  springs  or  seepages  in  the 
plateau  region  studied  has  a  fiow  of  more  than  a  few  gallons  an 
hour. 

It  is  freciuently  a  number  of  miles  from  one  spring,  ''tank"  or 
other  permanent  water  source  to  another. 

FIELD   WORK. 

In  the  summer  of  1921  and  1922,  Mr.  Moore,  with  a  party  of  four 
assistants,  was  assigned  by  the  United  States  Geological  Survey  to 
make  an  examination  of  a  portion  of  the  high  plateaus  of  southern 
Utah,  with  reference  to  coal  resources  and  possibilities  of  oil  and 
gas  development.  A  detailed  geological  map  of  approximately 
3,000  square  miles  was  made,  and  reconnoissance  study  of  a  very 
much  larger  area  was  completed.  The  region  mapped  comprises 
most  of  central  eastern  Garfield  county,  a  portion  of  central  Wayne 
county,  and  eastern  Kane  county,  Utah.  Although  the  primary 
purpose  of  this  work  had  to  do  with  possible  coal,  oil  and  gas  re- 
sources, special  attention  was  given  to  water  supply,  for  the  region 
is  a^emidesert.  Not  only  was  it  important  for  the  party  to  find 
water  for  camp  purposes,  but  the  possible  development  of  water 
supply  is  in  all  cases  the  most  important  consideration  in  the  utiliza- 
tion of  such  a  region.  During  the  course  of  this  work,  wherever 
water  insects  were  found,  and  where  it  was  possible  without  inter- 
ruption of  the  main  objects  of  the  work,  collections  were  made. 
Since  no  other  similar  collections  have  been  reported  from  this 
region,  and  since  the  distribution  of  water  insects  in  this  region  is 
both  of  biological  and  general  entomological  interest,  the  results 
are  here  presented.  Identification  and  special  notes  on  the  species 
found  are  the  work  of  Mr.  Hungerford. 

DESCRIPTION    OF    COLLECTIONS. 

Since  the  collections  of  water  insects  to  be  noted  below  were 
gathered  as  opportunity  offered  and  not  as  a  part  of  a  systematic 
faunal  survey,  the  data  are  perhaps  somewhat  fragmentary  and 
scattered.     However,  since  the  sources  of  water  supply  in  such 


416  The  University  Science  Bulletin. 

country  are  not  numerous,  and  since  it  is  necessary  for  the  traveler, 
whatever  his  mission,  to  seek  the  places  where  water  may  be  had, 
it  is  probable  that  a  considerable  number  of  the  places  where  water 
insects  might  occur  were  observed.  In  each  of  these  a  collection, 
representing  as  far  as  possible  all  of  the  species  present,  was  ob- 
tained. The  collections  include  mainly  those  from  springs  and 
"tanks."  It  is  possible  that  water  insects  could  have  been  found 
in  some  of  the  seepages  along  streams  or  in  the  permanent  streams, 
but  there  was  no  opportunity  to  make  special  search  for  these. 

SMALL  "tank,"  one  AND  ONE-HALF   MILES  ABOVE  MOUTH   OF   MULEY 
TWIST   CREEK,  EASTERN   GARFIELD   COUNTY,  UTAH. 

Muley  Twist  creek  is  an  intermittent  stream  whose  bed  is  dry 
the  greater  part  of  the  year.  It  occupies  a  very  deep  box  canyon, 
carved  in  massive  red  and  yellow  sandstone.  About  seventy-five 
feet  above  the  bottom  of  the  canyon,  on  its  sloping  west  sandstone 
wall,  was  found  at  one  point  about  one  and  one-half  miles  above 
the  mouth  of  the  creek,  a  little  "tank"  about  one  by  two  feet  in 
width  and  length  and  with  greatest  depth  of  approximately  one 
foot.  The  depression  was  filled  with  clear  water,  part  probably 
caught  in  a  recent  rain  and  part  derived  from  a  very  small  seep  in 
the  sandstone  above.  The  "tank"  is  probably  not  at  all  permanent. 
It  was  surrounded  by  bare  smooth  sandstone,  without  any  near-by 
plant  growth. 

In  this  little  basin  lived  a  group  of  seven  individuals  of  Arcto- 
corixa  abdominalis  Say,  without  other  insect  life. 

"tank"   about    two    MILES    ABOVE    MOUTH    OF    MULEY    TWIST    cllEEK, 
EASTERN   GARFIELD    COUNTY,    UTAH. 

On  the  east  side  of  the  Circle  Cliffs  and  along  the  Water  Pocket 
Fold  in  eastern  Garfield  county  are  a  number  of  "tanks"  which  con- 
tain water  in  all  but  the  driest  seasons.  From  one  of  these,  in  the 
bed  of  Muley  Twist  creek,  beneath  a  high  sandstone  cliff,  a  number 
of  water  bugs  were  taken.  The  pool  was  about  eight  feet  in 
diameter  and  three  feet  deep  in  one  part  at  the  time  of  our  visit  in 
August.  Two  or  three  large  cottonwoods  shade  the  pool,  but  there 
is  almost  no  other  vegetation  near  at  hand. 

In  this  little  basin  was  taken  one  dytiscid  beetle,  Rhanhis  bino- 
tatus  Harr.,  and  upon  the  surface  eight  specimens  of  Gerris  orba 
Stal,  all  winged,  and  one  pair  mating  (five  males  and  three  females). 
The  solitary  dytiscid  beetle  appears  to  be  a  new  record  for  Utah. 
Its  previous  capture  has  been  recorded  for  Wisconsin  and  Arizona. 


Moore  and  Hungerford:   Water  Insects.  417 

"tank"  on  COLORADO  RIVER  AT  WATER  POCKET  FOLD,  KANE  COUNTY. 

This  was  located  on  the  north  bank  of  the  Colorado  river  near  the 
mouth  of  a  small  tributary  canyon  at  Bennett's  oil  camp,  where 
Water  Pocket  Fold  crosses  the  canyon  of  the  Colorado  river,  about 
eight  miles  above  the  mouth  of  Escalante  river,  eastern  Kane 
county.  A  pool  of  clear  rain  water  filled  the  depression  in  bare 
sandstone,  about  four  feet  wide,  five  feet  long  and  two  and  one-half 
feet  deep;  no  vegetation  around  the  pool;  about  thirty  yards  from 
Colorado  river,  a  swift,  muddy  stream,  very  unlike  the  water  in  the 
"tank";  unknown  distance  to  adjacent  pools  of  clear  quiet  water; 
elevation  about  3,500  feet.    Here  were  taken: 

Fifteen  Arctocorixa  abdominalis  Say,  four  males,  eight  females 
and  three  nymphs.  This  represents  the  farthest  north  record  for 
the  species,  which  was  described  from  Mexico  and  reported  from 
Texas  and  California  by  Uhler. 

One  Notonecta  insulata  Kirby,  female. 

Two  Thermonectes  marmoratus  Hope,  one  male  and  one  female. 
The  capture  of  this  beautiful  dytiscid  appears  to  be  a  new  record  for 
the  state.  Leng's  catalogue  records  it  from  Arizona  and  Lower 
California. 

SPRING  OPPOSITE   OWL  CAVE,   HARRIS  WASH,  GARFIELD   COUNTY,   UTAH. 

About  twenty  miles  southeast  of  the  little  town  of  Escalante, 
eastern  Garfield  county,  Utah,  a  permanent  spring  is  found  in  the 
sandstone  wall  of  Harris  canyon,  which  is  tributary  to  Escalante 
river.  The  fiow  of  this  spring  is  not  large,  but  the  water  is  of  excel- 
lent quality,  and,  as  natural  shelter  is  conveniently  near  at  hand  in 
a  large  sandstone  caveYn,  Owl  Cave,  the  spot  is  a  camping  place  for 
most  of  the  few  travelers  who  pass  this  way.  The  water  from  the 
spring  irrigates  naturally  several  square  yards  of  protected  ground 
adjacent  to  the  spring.  A  beautiful  little  meadow,  therefore,  has 
been  formed,  through  which  the  water  flows  a  short  distance  before 
sinking  into  the  sand. 

This  spring  and  short  stream  support  a  populous  community  of 
aquatic  insects  and  other  life.  However,  only  two  species  of  water 
bugs  were  observed,  Gerris  remigis  Say  and  Trepobates  pictus  Uhl. 

SPRING  IN  WATER  POCKET  FOLD,  CIRCLE  CLIFFS,  GARFIELD  COUNTY, 

UTAH. 

One  of  the  very  few  water  sources  in  the  northeastern  side  of  the 
Circle  Cliffs  is  a  small  spring,  from  which  the  water  flows  a  few 
yards  before  disappearing  in  the  floor  of  a  canyon  tributary  to 


418  The  University  Science  Bulletin. 

Muley  Twist  creek.  The  adjacent  rocks,  chiefly  sandstone,  are 
nearly  bare  of  vegetation,  though  not  far  distant  are  numerous 
pinon,  scrub  cedar  and  sagebrush.  The  spring  varies  somewhat  in 
volume,  but  does  not  commonly  dry  up  in  the  spring  dry  season. 

On  the  surface  of  the  spring  and  the  little  stream  below  are 
numerous  representatives  of  the  species  Trepobates  pictus  and  Gerris 
remigis.  A  large  number  of  individuals  of  each  were  obtained,  in- 
cluding a  considerable  number  of  nymphs  belonging  to  the  first 
species.  The  Trepohates  were  more  numerous  than  the  Gerrids  at 
the  time  of  obser^'ation  in  August,  1921 — a  rather  unusual  condition 
as  compared  with  other  localities  inhabited  by  the  two  forms. 

SPRING  on  upper  HENRIEVILLE  CREEK,  SOUTH  CENTRAL  GARFIELD 

COUNTY,    UTAH. 

An  area  of  special  scenic  and  geologic  interest  in  southern  Utah 
is  Table  Cliff  and  the  surrounding  very  rough  district  in  south 
central  Garfield  county.  South  of  Table  Cliff,  on  one  of  the  branches 
of  upper  Henrieville  creek,  is  a  seepage  of  very  alkaline  water,  the 
only  source  available  for  camp  use  in  the  head  canyons  under  the 
cliff.    The  water  is  so  strongly  mineralized  as  to  be  hardly  potable. 

On  the  surface  of  the  short,  trickling  stream  below  the  seep  several 
water  bugs  were  taken.  A  few  Gerris  remigis  Say  and  a  considerable 
number  of  Micro  velia  are  the  inhabitants  of  this  apparently  in- 
hospitable haven.  The  Microvelia  species  of  this  region  has  not 
been  determined.    It  may  be  new. 

"tank"   NEAR  SOUTH   POINT   OF    KAIPAROWITS   PLATEAU,   EASTERN 

KANE   COUNTY,  UTAH. 

Most  of  the  observed  watering  places  in  southeastern  Utah  did  not 
contain  water  insects  at  the  time  of  visit  in  the  summer  of  1922.  Of 
four  springs  and  pools  on  the  top  of  Kaiparowits  plateau,  a  high 
tableland  which  extends  southeastward  toward  Colorado  river  from 
the  uplands  farther  north,  only  one  appeared  to  contain  any 
aquatics.  This  was  a  partially  rain-filled  "tank"  in  the  bottom  of 
one  of  the  narrow,  steep-sided  canyons  which  cut  the  plateau  sur- 
face. The  "tank"  was  carved  in  bare,  massive  sandstone,  without 
near-by  vegetation.  The  pool  was  about  six  feet  long  by  one  foot 
wide,  and  its  maximum  depth  a  little  over  one  foot.  In  this  pool 
were  several  Notonecta  insulata  Kirby,  but  no  other  water  bugs. 


MooHK  AM)  Ih  xgerford:   Water  Insects.  419 

LAST   CHANCE  CREEK,  EASTERN    KANE  COUNTY,   UTAH. 

In  ;i  pnrt  of  the  sandstone  canyon  of  Last  Chance,  about  twenty- 
five  miles  :il)()\-e  the  point  where  it  flows  into  Colorado  river,  a 
small  stream  Hows  more  or  less  permanently.  Other  parts  of  the 
canyon  contain  intermittent  flow,  but  in  most  cases,  except  for 
occasional  groups  of  Gerris  remigis,  did  not  contain  other  observed 
water  bugs.  At  the  place  described,  a  group  of  small  dytiscids,  a 
number  of  MicrovcUa  and  some  gerrids  were  collected. 

SPRING  ON   PARIA  RIVER,  ABOUT  FIFTEEN   MILES  BELOW   CANNONVILLE, 

KANE   COUNTY,   UTAH. 

Most  of  the  lower  Paria  river,  the  main  stream  in  south  central 
Kane  county,  Utah,  is  dry  the  greater  part  of  the  year.  The  river 
has  carved  a  very  deep  sandstone  canyon,  which  affords  the  only 
passageway  from  north  to  south  across  this  part  of  the  country.  At 
a  point  about  fifteen  miles  below  the  town  of  Cannonville,  the  last 
settlement  on  the  southward  route  to  Colorado  river,  at  Lee's  Ferry, 
about  seventy-five  miles  distant,  is  a  fine  spring.  The  flow  is  fairly 
large,  as  compared  with  other  springs  in  this  country,  and  the  water 
is  clear  and  not  alkaline. 

A  host  of  Gerris  remigis  and  Trepobates  pictus  were  found  on  the 
water  of  the  spring. 

LOWER  ROCK  CREEK,  EASTERN   KANE  COUNTY,  UTAH. 

Water  fl.ows  more  or  less  permanently  in  the  lower  part  of  Rock 
creek,  the  easternmost  of  the  longer,  deep  canyons  tributary  to  Colo- 
rado river  west  of  Kaiparowits  plateau.  The  canyon  of  Rock 
creek,  as  the  name  perhaps  suggests,  is  somewhat  unusual,  even 
among  the  great,  bare  rock  cliffs  and  canyons  of  the  plateau  coun- 
try. Its  walls  and  bottom  are  mostly  composed  of  naked  rock,  and 
sculpture  by  running  water  has  produced  an  almost  indescribably 
rough  topography. 

The  stream  at  the  camp  site  of  the  writer,  about  two  miles  above 
Colorado  river,  contained  numerous  Microvelia  and  a  few  gerrids. 
No  other  water  bugs  were  observed. 

NEAR  GOODRIDGE,    SAN    JUAN    COUNTY,   UTAH. 

In  a  small,  isolated  spring,  about  one  foot  in  diameter  and  a  few 
inches  deep,  were  taken  three  Notonecta  insulata  Kirby.  No  other 
water  insects  were  present.  This  tiny  basin  of  water  was  three  miles 
from  the  swift,  muddy  San  Juan  river,  and  there  were  no  other  near 
water  sources  known.    The  location  of  tiie  spot  is  at  the  west  side 


420  The  University  Science  Bulletin. 

of  Comb  Ridge,  where  the  road  from  Bluff  to  Goodridge  enters 
Comb  Wash.    Some  vegetation  was  growing  about  the  spring. 

To  the  above  Utah  collections  may  be  added  this  interesting  one 
from  northwestern  Colorado,  made  in  1920. 

junction  mountain,  YAMPA  river,  northwestern  COLORADO. 

In  the  deep  canyon  of  Yampa  river,  where  the  river  plunges 
through  Junction  mountain,  about  thirty-five  miles  east  of  the  con- 
fluence of  the  Yampa  and  the  Green,  in  northwestern  Colorado,  were 
taken  forty-four  Rhagovelia  distincta  Champ.,  all  apterous;  nine 
were  males,  four  nymphs,  and  the  remainder  females;  also  seven 
Trepobatopsis  trux  Bueno,  two  males  and  five  females,  described  by 
Mr.  Bueno  as  new. 

From  a  small,  sluggish  stream  at  west  side  of  Junction  mountain 
were  taken  an  interesting  series  of  large  gerrids.  These  vary  in 
color  from  russet  to  the  dark  color  typical  of  Gerris  remigis  Say. 
The  russet-colored  form  has  been  determined  by  Mr.  J.  R.  de  la 
Torre  Bueno  as  Gerris  orba  Stal.  Of  these  there  are  nineteen  speci- 
mens, nine  males  and  ten  females.  Thirteen  of  the  lot  are  apterous. 
G.  orba  Stal  has  been  listed  from  California,  Oregon  and  Nevada 
hitherto.  The  others,  forty-seven  of  them,  all  apterous,  are  of  a 
trifle  lighter  color  than  G.  remigis  Say,  but  obviously  are  the  same 
as  Bueno's  orba. 

In  an  attempt  to  find  differences  between  our  common  Gerris  re- 
migis Say  and  this  series  from  the  West,  all  of  the  Kansas  and  Colo- 
rado material  available  (a  series  from  the  eastern  margin  of  Kan- 
sas through  the  state  and  across  Colorado)  has  been  very  carefully 
studied.  The  Kansas  forms  are  darker  and  larger  as  a  series,  but  it 
does  not  seem  possible  at  the  present  time  to  fix  upon  any  structural 
characters  to  distinguish  these  forms. 

In  connection  with  the  problem  of  separating  the  striders,  it  may 
not  be  out  of  place  to  state  that  it  seems  to  have  been  overlooked 
that  Gerris  conformis  Uhl.  males  have  the  ventral  side  of  the  sixth 
abdominal  segment  singly  emarginate,  like  Limnoporus  rufoscutel- 
latus  Latr.  Indeed,  the  two  species  have,  in  addition  to  the  above 
genital  character,  another  distinctive  character  that  is  common  to 
both,  namely,  their  very  long  legs.  The  hind  femora  in  both  spe- 
cies greatly  surpass  the  tip  of  the  abdomen.  The  examination  of 
such  material  as  has  been  available  suggests  that  the  two  species 
commonly  accepted  as  L.  rujoscutellatus  and  Gerris  conformis  Uhl. 
may  be  separated  as  follows : 


Moore  AND  HuNGERFORD :  Water  Insects.  (^y    ^^/ 

A.  Antennal  segment  1  longer  than  2  plus  3.  Eyes  protuberant.  Middle  and 
hind  femora  about  equal  in  length.  Venter  of  sixth  abdominal  segment 
in  male  with  median  longitudinal  fossa.  The  seventh  with  a  longitudinal 
ventral  carina.  Gerris  conjormis  Uhl. 

AA.  Antennal  segment  1  shorter  than  2  plus  3.  Eyes  not  protuberant.  Middle 
femora  shorter  than  hind  femora.  Venter  of  sixth  abdominal  segment  in 
male  without  fossa  and  seventh  segment  not  carinate. 

Liinnoporus  rufoscutellatus  Latr. 

In  addition  it  may  be  stated  that  L.  rufoscutellatus  Latr.  is  a 
more  slender  species,  and  that  it  is  usually  of  a  russet  color.  Gerris 
conjormis  Uhl.  has  the  anterior  part  of  the  pronotum  bearing  two 
prominent  papillae  and  well-elevated  callosities. 

It  would  seem  somewhat  of  a  question  whether  these  two  species, 
having  in  common  genital  characters  and  exceptionally  long  femora, 
can  be  placed  in  separate  genera  upon  the  antennal  character  alone. 
Structurally  they  are  more  nearly  congeneric  than  G.  remigis  Say 
and  G.  conjormis  Uhl.  The  latter  belongs  with  L.  rujoscutellatus 
Latr.,  and  the  generic  characters  of  Limnoporiis  should  be  revised. 

resume. 

In  brief  resume  of  the  above  collecting  notes,  it  may  be  said  that 
the  southern  Utah  desert  affords  extremely  interesting  problems  for 
the  student  of  water  life.  The  light' annual  rainfall,  the  isolation  of 
the  pools  and  the  nature  of  the  water  they  contain,  lend  interest  to 
the  study  of  the  resident  population  of  the  waters.  The  collections 
here  reported  represent  the  complete  insect  population  of  the  various 
pools  surveyed.  One  isolated  little  spring-fed  pool,  the  size  of  a 
washbasin,  contained  three  Notonecta  insulata  Kirby,  and  no  other 
insects.  Another  little  rain-filled  "tank,"  less  than  two  feet  in  diam- 
eter and  a  foot  deep,  located  high  up  on  the  sloping  wall  of  a  canyon, 
contained  seven  water  boatmen,  Arctocorixa  abdominalis  Say,  and 
not  another  insect.  The  scant  annual  rainfall  of  from  five  to  ten 
inches,  with  the  exceptionally  light  fall  in  April,  May  and  June, 
works  against  a  large  population  of  aquatics.  The  water  bugs 
thus  utilize  every  available  supply  of  water.  The  pools  contained 
the  corixids,  notonectids  and  dytiscids,  while  the  striders  were  found 
on  the  springs  more  commonly  than  upon  the  "tanks."  The  pres- 
ence of  winged  species  in  places  remote  from  permanent  water  is 
readily  understood.  The  finding  of  eight  Gerris  orba  Stal,  all 
winged,  upon  a  rain-filled  "tank"  and  forty-seven  specimens  of  the 
same  species  along  the  Yampa  river,  all  apterous,  is  worthy  of  note.* 

*  An  interesting  note  on  the  factor  regulating  wing  development  is  suggested  by  collect- 
ing notes  on  Microvelia  macgregori  Kirk,  in  New  Zealand  by  Mr.  J.  G.  Myers,  in  the  New 
Zealand  Journal  of  Science  and  Technology,  vol.  V,  pp.  6  and  7.  He  writes  that  his  col- 
lections of  this  insect  in  the  lowlands  have  been  almost  entirely  apterous  forms,  while  at  an 
altitude  of  4,500  feet  every  capture  was  winged. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 

Vol.  XIV,  No.  18— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  18.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

The  Xepid.e  in  North  America  North  of  Mexico, 

H.  B.  Hungerford. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE  KAN. 


Entered  at  the  po.st  office  in  Lawrence  as  second-class  matter. 

9-4522 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  18. 


I 


The  Nepida3  of  North  America. 

(Further  Studies  in  Aquatic  Hemiptera.) 

BY  H.   B.   HUNGERFORD. 

ACKNOWLEDGEMENTS. 

WISH  herewith  to  acknowledge  my  indebtedness  to  those  who 
have  rendered  assistance  in  the  preparation  of  this  paper,  which 
had  its  beginning  several  years  ago.  I  am  under  especial  obligation 
to  Dr.  L.  O.  Howard,  Dr.  J.  M.  Aldrich,  and  Mr.  W.  L.  McAtee  for 
the  opportunity  they  gave  me  to  study  in  the  United  States  National 
Museum,  where  are  located  some  of  the  types  of  Doctor  Montandon 
and  the  Kirkaldy  and  Uhler  collections  of  these  insects.  In  addi- 
tion to  the  material  at  the  United  States  National  Museum,  I  have 
been  permitted  to  examine  the  collections  of  Mr.  J.  R.  de  la  Torre 
Bueno,  of  Dr.  Carl  J.  Drake  and  have  had,  beside  our  own  Univer- 
sity collections,  abundant  material  so  generously  supplied  by  Mr. 
W.  E.  Hoffmann  and  Doctor  Knight,  of  the  University  of  Minne- 
sota, and  by  Mrs.  Grace  Wiley  from  collections  in  Texas.  Profes- 
sor Brimley,  of  Raleigh,  N.  C,  also  supplied  me  with  two  species 
for  study.  Miss  Kathleen  Doering  made  most  of  the  drawings  and 
Mr.  P.  A.  Readio  and  Mr.  Raymond  Beamer  helped  me  with  the 
photographs. 

INTRODUCTION. 

The  subtle  characters  and  elusive  specific  differences  between  the 
various  species  of  the  cryptocerate  groups  of  the  Hemiptera  have 
made  it  difficult  to  fix  specific  limits.  The  students  of  these  groups 
have  resorted,  therefore,  to  comparisons  that  are  beyond  the  compre- 
hension of  the  general  systematist  endeavoring  to  identify  his  col- 
lection. Such  comparative  notes  become  intelligible  only  after  a 
prolonged  study  of  the  insect  group  concerned,  and  this  condition 

(425) 


426  The  University  Science  Bulletin. 

has  worked  against  there  being  a  very  large  number  of  students 
who  have  found  the  time  and  inclination  to  identify  the  species  in 
such  families  as  the  Notonectidse,  Nepida?  and  Corixidse,  for  in- 
stance. 

The  difhculty  encountered  by  the  writer  in  sorting  and  naming 
the  aciuatic  bugs  in  the  course  of  his  biological  studies  led  to  a  thor- 
ough investigation  for  diagnostic  characters  of  more  demonstrable 
kind  than  those  employed  in  the  literature.  It  has  been  the  en- 
deavor, therefore,  to  find  and  figure  characters  of  sucli  definiteness 
that  anyone  with  even  fair  training  in  close  work  can  be  certain  of 
the  species  he  is  trying  to  identify. 

The  first  report  upon  this  work  was  given  under  the  title,  "The 
Male  Genitalia  as  Characters  of  Specific  Value  in  Certain  Crypto- 
cerata,"  which  appeared  in  volume  XI  of  the  Kansas  University 
Science  Bulletin,  December,  1919.  In  this  the  writer  called  at- 
tention to  these  characters  in  the  Naucoridse,  Saldidae  (not  a  crypto- 
cerate,  of  course),  Gelastocoridae,  Corixidse  and  Notonectidse.  The 
treatment  of  the  genus  Notonecta  was  sufficiently  complete  to  be  of 
value  to  the  systematist,  because  the  male  genital  capsules  of  all  the 
North  American  species  were  figured  or  described,  save  two,  A\  iM- 
eri  Kirk,  and  N.  montezuma  Kirk.,  the  former  readily  recognizable 
by  the  description,  and  the  latter  exceedingly  rare,  repre.>:nted,  so 
far  as  indicated  in  the  literature,  by  two  specimens  in  the  Hope 
museum,  carefully  studied  and  figured  by  Champion.  (Figure  8, 
plate  XXXI,  Science  Bulletin  XI,  figures  Notonecta  howarclii  Bueno, 
and  fig.  5,  plate  XXXI,  Science  Bulletin  XI,  figures  Notonecta 
shooteriiVhl.  The  one  there  named  N.  lutea  is,  of  course,  N.  borealis 
Hussey  and  Bueno.) 

Studies  were  made  upon  the  Nepidse  and  Corixidae,  but  were  not 
reported  in  the  first  paper,  and  there  was  no  intention,  until  recently, 
of  publishing  upon  the  Nepidse.  However,  four  species  of  Ranatra 
have  been  taken  in  Kansas  waters  as  a  result  of  recent  collecting, 
all  new  records  for  the  state.  The  task  of  naming  these  water  scor- 
pions involves  problems  in  nomenclature  and  the  authenticity  of 
species — matters  concerning  which  there  is  a  difference  of  opinion 
among  our  best  authorities. 

Mr.  J.  R.  de  la  Torre  Bueno  has  believed  that  Ranatra  nigra  Her- 
rich  Schaffer  1853,  is  a  synonym  of  Ranatra  fusca  Palisot  Beauvois 
1805.  The  insect  which  he  calls  Ranatra  fmca  is  large  with  very 
prominent  eyes,  slender  anterior  legs,  and  a  deep,  broad  prosternal 
groove — a  very  characteristic  insect.     With  this  interpretation  of 


Hlxgekfurd;    Xki'ii).!-:  im  Amerra.  427 

Palisofs  species,  he  described  a  small,  compactly  built  insect  with 
\ei'v  broad  front  femora,  as  Ranatra  kirkaldyi. 

Dr.  A.  L.  Montandon  (Bui.  Soc.  Sci.  Bucharest,  XIX,  1910)  con- 
siders Ranatra  kirkaldyi  Bueno  a  synonym  of  Ranatra  fusca  P.  B., 
and  concludes  that  Ranatra  nigra  H.  S.  is  a  good  species,  although 
he  has  seen  no  American  specimen  that  fits  the  description.  Van 
Duzee  follows  Montandon. 

In  a  recent  issue  of  the  Entomological  News  (vol.  XXXII,  p. 
273,  1921),  under  the  title  "New  Records  of  Aquatic  Hemiptera  for 
the  United  States  with  Description  of  New  Species,"  Mr.  Torre 
Bueno  again  defends  his  Ranatra  kirkaldyi  and  comments  on  Ra- 
luitra  fusca  P.  B.,  describing  it  so  closely  that  there  can  be  no  mis- 
taking the  insect,  and  stating  that  there  is  a  specimen  of  this  species 
in  the  United  States  National  Museum  labeled  Ranatra  fusca  P.  B., 
by  Doctor  Montandon.    This  statement  is  indeed  true. 

Through  the  kindness  of  Dr.  L.  O.  Howard,  Doctor  Aldrich  and 
W.  L.  McAtee;  the  writer  had  the  pleasure  of  spending  some  time  at 
the  United  States  National  Museum  studying  the  aquatic  Hemip- 
tera. He  had  the  opportunity,  therefore,  of  examining  the  material 
determined  by  Doctor  Montandon,  as  well  as  the  types  of  Ranatra 
protema  Montd.,  Ranatra  brevicollis  Montd.  and  Curicta  hoivai'dii 
Montd.  He  found  two  distinct  species  identified  by  Doctor  Mon- 
tandon as  R.  fusca  P.  B.  One  of  them  is  Bueno's  Ranatra  fusca; 
the  other,  cjuite  a  different  species,  but  not  Bueno's  R.  kirkaldyi. 

After  examining  much  material  in  the  family  Nepidse,  and  very 
carefully  studying  and  weighing  original  descriptions  and  compara- 
tive notes,  the  writer  has  been  forced,  in  spite  of  his  reluctance,  to 
a  new  interpretation  of  the  old  species.  The  facts  which  led  to  the 
change  in  nomenclature  are  presented  under  the  species  concerned. 
All  the  species  are  figured  in  such  detail  that  there  need  be  no  ques- 
tion or  uncertainty  as  to  designation,  and  these  studies  should  aid 
in  arriving  at  a  solution  of  the  matters  in  controversy. 

Tlie  family  Nepidse  is  represented  in  America,  north  of  Mexico, 
by  three  genera,  Nepa,  Curicta  and  Ranatra,  which  may  be  sepa- 
rated as  follows: 

A.   Body  broadly  oval  and  flat.  Nepa. 

AA.   Body  elongate. 

B.    Prothorax  a  little  broader  than  head,  body  elongate  oval. 

Curicta, 

BB.    Prothorax  narrower  than  head,  body  ver>^  elongate.        Ranatra. 

Genus  Sepa,  see  p.   428;    genus  Curicata,  see  p.  429;    genus  Ranatra.  see  p.   433. 


428  The  University  Science  Bulletin. 

Genus  Nepa  Linnaeus  1758. 
We  have  one  species  in  the  genus  Nepa:  N.  apiculata  Uhl. 

Nepa  apiculata  Uhler. 

Uhler,  P.  R.,  in  T.  W.  Harris'  Insects  Injurious  to  Vegetation,  3d  ed.,  p.  12,  plate  1,  fig. 
1  f   1862. 

This  species  was  first  made  known  to  science  through  a  picture  of 
it  which  appeared  in  the  third  edition  of  Harris'  "Insects  Injurious 
to  Vegetation."  In  this  edition  Doctor  Uhler  added  notes  on  the 
Hemiptera.  Then  in  1878,  Uhler,  in  his  "Notices  of  the  Hemiptera 
Heteroptera  in  the  Collection  of  the  Late  T.  W.  Harris"  (Proceed- 
ings of  the  Boston  Society  of  Natural  History,  vol.  XIX,  Pt.  IV, 
1878),  records  the  presence  of  "No.  26,  Harris  Collection,  Nepa  apic- 
ulata Say  MS.,  under  stones  near  water,  May  15,  1826,"  and  adds 
the  following  descriptive  note:  "The  principal  differences  between 
our  species  and  the  European  one  consist  in  the  color  of  the  tergum, 
which  is  red  in  the  latter,  fuscous  in  ours ;  and  the  length  of  the  api- 
cal tubes,  which  in  ours  are  stouter  and  shorter."  In  commenting 
on  these  comparisons,  Montandon  (Bui.  §oc.  Sci.  Bucharest,  VIII, 
1898)  says  that  Uhler,  like  Stal  and  Ferrari,  attached  too  much  im- 
portance to  color,  and  gives  illustrations  of  the  variability  of  Nepa 
apiculata  Uhl.  and  of  Nepa  cinerea  L.  to  prove  the  danger  of  con- 
sidering color  of  specific  significance.  He  then  states  that  the  Amer- 
ican species  is  a  little  more  transverse  across  the  thorax,  the  abdo- 
men proportionately  larger  in  the  rear,  and  the  respiratory  tubes 
shorter  than  in  the  European  species.  Again  in  volume  XVIII  of 
the  same  periodical,  under  the  title,  "Hydrocorises  de  I'Ame'rique  du 
Nord,  Notes  et  Descriptions  d'Especes  Nouvelles,"  Doctor  Mon- 
tandon adds  that  in  "Nepa  apiculata  Harris  1862,  Uhler  1847,* 
.  .  .  the  superior  part  of  the  head  is  less  boldly  carinate,  espe- 
cially on  the  vertex,  which  is  generally  almost  smooth,  cjuitc  feebly 
convex." 

These  differences  between  the  European  Nepa  cinerea  L.  and  the 
American  Nepa  apiculata  Uhl.  do  exist,  as  an  examination  of  the 
photographs  on  plate  LI  will  show.  However,  without  both  species 
for  study,  one  would  be  compelled  to  remain  in  doubt  or  name  the 
species  from  its  geographical  distribution.  Since  there  do  occur 
marked  structural  differences,  it  is  well  to  note  them.  The  antennae, 
for  instance,  are  very  different,  as  an  examination  of  figures  5  and 

*Probably  a  typographical  error,  because  Uhler  was  born  in  1835.  Van  Duzee  omits  the 
reference:  Montandon,  Bui.  Soc.  Sci.  Bucharest,  XVIII,  p.  180,  from  the  list  under  jVepa 
apiciblata  Uhl.  in  his  catalogue  of  1917.  It  is  interesting  to  note  that  Nepa  apiculata  was  a 
Say  manuscript  species. 


Hungerford:    Nepid^  in  America.  429 

6,  on  plate  XLVII,  will  show.  The  penultimate  segment  of  the 
European  form  has  a  lateral  prolongation  which  gives  the  antenna 
a  branched  appearance.  The  Nepa  apiculafa  lacks  this  entirely. 
The  male  genital  capsules  are  also  unlike.  The  considerable  differ- 
ence between  these  and  between  the  claspers  is  indicated  on  plate 
XLVII,  figures  1,  2,  3  and  4. 

The  nearest  approach  to  a  formal  description  of  the  American 
species  is  given  by  Uhler  in  the  Riverside  Natural  History,  vol.  II, 
p.  253,  1884.    It  follows: 

Color  (lull  fuscous  gray,  with  the  base  of  the  abdomen  above  more  or  less 
tinged  with  reddish.  It  is  of  an  elliptical  form,  blunt  in  front,  with  a  ridged 
middle  line  on  the  vertex,  and  with  three  short  raised  lines  on  the  prothorax. 
each  side  of  a  longer  one  on  the  middle.  The  surface  and  margins  of  the 
thorax  and  head  are  roughly  granulated,  while  these,  tog.ether  with  the  scu- 
tellum  and  corium,  are  rough  and  closely  covered  with  stiff,  short  pile.  The 
anterior  femora  have  no  teeth  on  the  inner  angle,  but  instead  there  is  a  prom- 
inent elbow,  forming  a  wide  expansion  for  the  sides  of  the  deep  gutter.  The 
wings  are  smoke  brown,  with  darker  veins.  This  species  closely  resembles  the 
European  one,  and  measures  about  twc^-thirds  of  an  inch  to  the  end  of  the 
abdomen;  while  the  respiratory  tubes  are  a  little  more  than  one-fourth  of  an 
inch  in  length. 

Montandon  has  shown  the  differences  between  our  species  and 
the  European  A''.  Cinerea  Linn. 

The  United  States  National  Museum  has  specimens  from  Massa- 
chusetts, Pennsylvania,  Maryland  and  Illinois.  I  have  seen  speci- 
mens from  New  York,  Minnesota  and  Wisconsin. 

Genus  Curicta  Stal,  1861. 
These  insects  are  very  interesting  because  they  are  intermediate 
between  the  broad,  flat  Nepa  and  the  very  slender  Ranatra.  The 
Americas  can  boast  of  several  species  in  this  genus,  but  for  North 
America,  north  of  Mexico,  there  has  previously  been  recorded  only 
one,  and  of  it  but  a  single  specimen,  taken  at  Victoria,  Tex.,  just 
within  our  limits.  From  this  specimen  Doctor  Montandon  drew  up 
the  description  of  Curicta  howardii.  It  is  a  pleasure,  therefore,  to 
record  the  capture  of  other  specimens  of  this  species  and  to  add  a 
second  species  which  has  not  been  described.  This  second  species 
has  been  the  subject  of  life-history  studies  by  Mrs.  Grace  Wiley, 
and  is  reported  further  on  in  this  bulletin. 

KEY  TO   SPECIES. 

A.   Lateral  prolongation  of  penultimate  segment  of  antenna  very  short  (see 
fig.  8,  pi.  XLVII).  C.  howardii  Montd. 

AA.   Lateral  prolongation  of  penultimate  segment  of  antenna  very  long  (see 
fig.  7,  pi.  XLVII).  C.  drakei  sp.  new. 


430  The  University  Science  Bulletin. 

.   Curicta  howardii  Montandon,  1910. 

Montandon,  A.  L.,  Bui.  Soc.  Sci.  Bucharest,  XVIII,  p.  181;   1910. 

The  original  description  is  in  French.  The  writer's  free  trans- 
lation follows: 

Elongate  oval  in  form,  visibly  attenuate  in  front  and  rear,  lateral  margins 
not  subparallel,  the  greatest  width  toward  the  posterior  third.  Head  quite 
enlarged,  although  a  little  narrower  than  the  front  part  of  the  pronotjuii,  as 
long  as  wide,  including  the  eyes,  longitudinally  carinate  throughout  its  length, 
the  carina  more  obtuse  on  the  posterior  interocular  portion.  Interocular 
space  more  than  three  times  as  wide  as  the  diameter  of  the  eye.  Eyes  small, 
globular,  anterior  part  of  head  triangular,  exceeding  the  anterior  level  of  the 
eyes  by  a  length  ecjual  to  its  width  between  the  eyes  in  front. 

Pronotum  distinctly  longer  than  its  width  behind,  lateral  edges  subparallel 
on  their  anterior  three-fifths,  quite  strongly  widened  on  their  posterior  two- 
fifths;  with  four  obtuse  longitudinal  carinse,  little  accentuated  and  sub- 
parallel,  two  each  side  of  the  anterior  part,  the  posterior  part  with  two 
oblique  carinae  arising  from  the  anterior  median  carina  and  quite  divergent 
behind.  The  anterior  depression '  of  the  pronotum  broadly  semicircular,  the 
anterior  angles  quite  narrowed,  subacute. 

Scutellum  with  three  longitudinal  carinas,  the  median  continuing  quite 
plainly  clear  to  the  apex  of  the  scutellum.  The  two  laterals  slightly  diverging 
behind,  ^•anishing  on  the  middle  of  the  sides  of  the  scutellum,  which  are 
slightly  sinuate  before  the  tip  or  end. 

Coria  insensibly  and  gradually  widened  behind  on  their  basal  halves,  at- 
taining their  greatest  width  behind  the  middle  and  narrowing  thereafter; 
membrane  well  developed,  regularly  subrounded  at  the  extremity.  Com- 
missure of  the  clavus  almost  twice  as  long  as  the  scutellum. 

Appendages  short,  quite  robust  toward  the  base,  attenuated  thereafter, 
about  half  the  length  of  the  abdomen. 

Anterior  femora  quite  robust,  as  long  as  the  pronotum  on  its  lateral  edgeSj 
with  a  single  median  tooth  easily  visible  on  the  inner  edge  of  the  groove  where 
the  folded-up  tibia  is  lodged,  this  tooth  distinctly  closer  to  the  base  than  to 
the  apex  of  the  femur;  the  external  side  of  the  groove  also  appears  denticu- 
late, as  if  notched  on  the  basal  third  of  the  femur.  Neither  teeth  nor  sinuosi- 
ties toward  the  apex  of  the  femur. 

Anterior  coxae  half  the  length  of  their  femora.  Anterior  tibia  quite  long, 
blackish,  with  a  pale  annulation  toward  the  base,  and  the  apical  third  like- 
wise pale;  the  extremity  of  the  tarsi  come  to  the  basal  third  of  the  femora 
when  the  tibia  is  folded  back  against  the  latter. 

Intermediate  and  posterior  legs  short,  the  ends  of  the  posterior  femora, 
which  are  a  little  shorter  than  their  tibiae,  do  not  reach  the  suture  of  the  last 
abdominal  segment.  Intermediate  and  posterior  tarsi  with  their  claws  less 
than  half  as  long  as  their  tibiae. 

Median  longitudinal  part  of  the  prosternum  slightly  saddle-shaped,  pro- 
jecting in  all  its  width,  more  elevated  than  the  lateral  pieces,  a  little  flat- 
tened and  traversed  its  whole  length  by  a  fine  median  groove;  very  obtusely 
tuberculate  in  its  anterior  part.  A  greater  space  between  the  intermediate 
coxae  than  between  the  anterior  or  posterior  coxae. 

Length,  19  mm.;  maximum  width  a  little  behind  the  middle  of  the  corium, 


Huxgkkfokd:    Nkpid.?<:  in  America.  431 

4.5  mm.;  at  bn^^v  ol'  I'lonotum.  3.8  mm.;  length  of  appendages,  7.7  mm.,  Vic- 
toria. Tox.    A  single  specimen,  United  States  National  Museum,  Washington. 

Tt)  the  above  description  Doctor  Montandon  adds  the  following 
helpful  coniparative  notes: 

This  species  is  intermediate  in  size  between  C.  volxemi  Montd.  and  C. 
Scorpio  Stal  (.=>nonta)idom  Martin).  It  differs  from  C.  voh-cini  Montd.  by 
the  iironotiun  being  sensibly  narrowed  in  front,  while  in  the  latter  it  is  almost 
as  wide  in  front  as  behind.  In  this  character  it  approaches  more  closely  to  C. 
Scorpio  Stal,  which  also  has  the  pronotum  quite  narrowed  in  front,  with  the 
same  right-angled  anterior  angles  almost  sharp,  but  its  anterior  tibife  are, 
however,  slightly  more  elongated  than  in  this  latter  species;  that  is  to  say, 
much  more  than  in  C.  volxemi  Montd.  In  the  character  of  the  median  tooth 
of  the  anterior  femur  being  situated  closer  to  the  base  than  the  extremity, 
howe\-er.  the  species  approaches  C.  scorpio  Stal,  but  it  is  plainly  separated 
from  the  latter  by  its  head  being  longitudinally  carinate  throughout  its  entire 
length,  while  the  head  is  simply  convex  between  the  eyes  in  C.  scorpio  Stal 
and  almost  plain  in  C.  volxemi  Montd. 

Furthermore,  in  C.  scorpio  the  scutellum  is  not  carinate;  the  longitudinal 
grooves  of  the  pronotum  are  also  much  less  emphasized  and  the  anterior 
tibiae  are  more  largelj*  pale,  darker  only  toward  the  base. 

This  is  the  first  species  of  the  genus  found  in  the  United  States.  It  is 
to  b(>  ]iresumed,  however,  that  others  may  occur  in  the  Southern  states 
neighboring  to  Mexico,  where  are  foimd  the  two  species  to  which  I  have 
just  compared  it. 

I  take  pleasure  in  dedicating  this  to  Mr.  L.  0.  Howard,  as  an  indeed 
feeble  expression   of  my  sincere   gratitude. 

In  the  collection  of  Prof.  H.  G.  Barber  are  three  males  from 
Huachuca  mountains,  Arizona,  taken  in  1899.  They  have  been 
compared  with  the  type  in  the  United  States  National  Museum 
by  Dr.  Carl  Drake,  through  the  kindness  of  whom  I  have  been 
privileged  to  study  the  structural  details  of  these  interesting  in- 
sects. The  United  States  National  Museum  has  a  specimen  taken 
by  D.  C.  Van  Dine  at  Camp  Travis,  Tex.,  March  30,  1918. 

The  species  is  a  compactly  built  creature.  The  head  is  set 
deeply  into  the  prothorax,  the  anterior  lateral  lobes  of  which  are 
conspicuously  swollen  and  incurved  on  their  anterior  tips  to  em- 
brace a  portion  of  the  eyes.  The  front  of  the  head  appears  almost 
truncate,  due  to  the  tylus  and  juga  being  about  equal.  The 
antennae  are  small  and  the  lateral  prolongation  of  the  penultimate 
segment  short  (see  fig.  8,  pi.  XLVII).  The  pronotum  has  a  broad, 
prominent  elevation  throughout  its  length.  On  the  anterior  half 
of  this  there  is  a  median  longitudinal  depression.  The  genital 
capsule  and  its  claspers  are  somewhat  different  from  those  of  the 
species  described  below  (see  figs.  7  and  8,  pi.  XLV) ,  but  the  specific 
differences  are  not  great. 


432  The  University  Science  Bulletin. 

Curicta  drakei  sp.  new. 

Size.  Length  from  16  mm.,  in  case  of  smallest  male,  to  22  mm.,  in  largest 
female — not  counting  the  respiratory  filaments,  which  are  from  7  to  8  mm. 
long.  The  width  of  the  head,  including  the  eyes,  from  2  mm.  to  2.25  mm.; 
the  greatest  width  of  anterior  part  of  thorax,  2.5  mm.  to  2.8  mm.;  the 
greatest  width  of  posterior  portion  of  thorax,  3  mm.  to  4  mm.;  the  greatest 
width  of  abdomen,  3.6  mm.  to  4.75  mm.;  the  length  of  pronotum  measvu-ed 
along  the  dorsal  median  line,  3,  mm.  to  3.6  mm. 

Color.  Obscured  by  incrustations,  which  color  it  from  gray  to  black. 
The  cleaned  insect  is  yellowish  to  dark  testaceous,  the  dorsum  of  abdomen 
red. 

Shape.  Relatively  slender;  greatest  width  of  thorax  is  to  length  of  body 
approximately  as  1  is  to  5.4.     Sides  of  thorax  and  abdomen  nearly  parallel. 

Structural  characteristics.  The  head  wider  across  the  eyes  than  distance 
from  caudal  margin  of  head  to  tip  of  lora.  The  length  before  the  eyes  greater 
than  that  behind  them.  Ej^es  small,  globular.  Tylus  longer  than  juga.  The 
head  longitudinally  carinate. 

Penultimate  segment  of  antennae  with  long,  slender  lateral  prolongation 
(see  fig.  7,  pi.  XLVII).  The  pronotum  with  anterior  lateral  lobes  not  greatly 
swollen;  the  lateral  margins,  therefore,  not  greatl.y  curved.  Three  longitudinal 
grooves  on  disc  of  pronotum,  the  median  one  broad  and  straight,  dividing 
the  median  longitudinal  elevation  into  two  parallel  carinse,  laterad  of  which 
are  the  deep  lunate  grooves  terminating  caiidolaterally  in  deep  depressions. 
The  posterior  enlarged  portion  of  pronotum  bears  four  more  or  less  distinct 
longitudinal  carinae,  which  in  some  specimens  appear  to  arise  as  bifurcations 
of  the  two  prominent  carinse  before  them,  and  again  as  independent  eleva- 
tions. Scutellum  tricarinate,  the  median  carina  more  prominent  on  posterior 
half,  the  lateral  carinse  slightly  curved  and  terminating  about  the  middle  of 
the  lateral  margins  of  the  scutellum,  these  elevations  and  declivities  often 
accentuated  or  obscured  by  incrustations.  Membrane  of  wings  well  de- 
veloped, as  long  as  the  elytral  suture,  reticulate,  the  apex  of  the  wing  cover- 
ing the  basal  two-fifths  of  the  genital  segment.  Operculum  of  male  genital 
segment  semicylindrical  and  slightly  constricted  before  the  apex,  which  is 
bluntly  pointed  and  faintly  carinrtte.  The  metasternal  plate  short,  caudal 
margin  nearly  straight,  the  posterolateral  prolongations  attaining  less  than 
half  the  length  of  the  coxae  and  exposing  a  large  elevated  area  (as  long  as 
wide)  of  sternite  behind  it.  Front  coxa  and  femiu-  stout,  coxa  one-half  length 
of  femur,  which  bears  its  rather  well-developed  tooth  nearer  the  base  than  the 
apex.  This  tooth  is  located  on  the  inner  edge,  two-fifths  of  the  distance  from 
the  trochanter  to  the  apex  of  the  femur.  Tibia  one-half  as  long  as  femur, 
the  apex  of  the  short  tarsal  segment  barely  attaining  the  middle  of  the  tooth 
when  tibia  is  flexed.  Mesothoracic  femur  one-fourth  longer  than  the  tibia, 
which  is  two  and  one-half  times  as  long  as  the  tarsus  without  the  claws. 
Metathoracic  femiu'  and  tibia  of  equal  length,  the  tibia  about  three'  and  one- 
half  times  the  length  of  tarsus  without  the  claws,  which  are  one-third  the 
length  of  the  tarsus. 

Notes.  The  above  species  is  described  from  a  series  of  20  speci- 
mens, 16  of  which  were  collected  by  Mrs.  Grace  Wiley  in  Colorado 


Hl\ni;i:kfori):    Xepid.k  in  Amkkica.  433 

county,  Texas.  Three  were  taken  in  New  Orleans,  La.,  and  belong 
to  Dr.  Carl  Drake;  and  one  from  tlie  state  of  Coloma,  Mexico,  is  in 
the  United  States  National  Museum.  The  species  is  named  in  honor 
of  Dr.  Carl  Drake,  who  believed  the  species  to  be  new  as  long  ago 
as  1916.  but  who  nc\-er  found  time  to  study  the  matter  thoroughly. 

Cvricta  drakeij  while  measuring  as  long  as  C.  houmrdii,  is  a  much 
more  slender  species,  and  appears  smaller.  Indeed,  comparing  the 
males  of  the  two  species,  there  is  a  considerable  difference  in  size 
and  shape.  The  anterior  lateral  lobes  of  the  pronotum  are  much 
more  prominent  in  C.  houmrdii,  the  sides  of  prothorax,  therefore, 
more  curved.  The  median  longitudinal  fossa  in  C.  howardii  is  con- 
fined to  anterior  part  of  the  median  elevation.  The  tylus  is  longer 
in  C.  drakei,  the  antennse  are  differently  formed  (see  figures  7  and  8 
on  plate  XLVII),  and  the  mietasternal  plate  is  smaller.  This  new 
species  differs  from  C.  voire tni  Montd.  from  Mexico  by  its  smaller 
size  (C  volxemi  is  241^  mm.  long  without  appendages),  by  the  tooth 
on  the  front  femur  being  conspicuously  nearer  base  than  apex,  while 
Doctor  ]\Iontandon's  figure  of  his  species  shows  the  tooth  in  the 
middle,  or  nearer  the  apex.  It  further  differs  from  C.  volxemi  by 
having  the  head  longitudinally  carinate  and  the  scutellum  tricari- 
nate.  The  thorax  is  relatively  shorter  than  in  C.  volxemi,  which 
has  a  thorax  twice  as  long  as  wide.  The  front  coxae  of  C.  drakei  are 
much  shorter. 

It  differs  from  C.  scorpio  Stal,  which,  according  to  Montandon, 
lacks  the  carina  on  head  and  scutellum  and  has  the  sides  of  the 
thorax  much  as  in  C.  howardii  Montd.  The  longitudinal  grooves  of 
the  thorax  of  C.  scorpio  Stal  are  less  emphasized  than  in  C.  howardii, 
which  in  turn  has  them  less  emphazised  than  C.  drakei.  All  of  these 
points  separate  C.  drakei  from  C.  scorpio  Stal,  and  are  based  upon 
Doctor  Montandon's  remarks  upon  the  latter  species.  Champion, 
in  his  "Biologia  Central!  Americana,"  places  Nepoidea  montandoni 
Martin  as  a  synonym  of  C.  scorpio  Stal.  Joanny  Martin,  under  the 
title  "Descriptions  d'Especes  Nouvelies  de  Nepidse  (Hem.),"  in  Bul- 
letin de  la  Societe  Entomologique  de  France,  1898,  pages  66-68,  de- 
scribes on  page  68  his  A^.  montandoni,  and  figures  the  head  and 
thorax  on  the  previous  page.  C.  drakei  is  unlike  this  species.  Stal's 
description  of  C.  scorpio  is  wholly  inadequate. 

Genus  Ranatra  Fabricius  1790. 
This  genus  is  characterized  by  its  very  elongate,  slender  form. 
It  is  the  dominant  genus  of  the  family  Nepidse  in  North  America. 
Doctor  Horvath,  in  his  paper,  "Les  Relations  entre  les  Faunes 


434  The  University  Science  Bulletin. 

Hemipterologiques  de  I'Europe  et  de  I'Amerique  du  Nord"  (Pro- 
ceedings of  the  Seventh  International  Zoological  Congress,  1907), 
lists  the  genus  Ranatra  as  belonging  to  the  oriental  fauna.  Since 
that  time  Doctor  Montandon  has  added  several  names  to  our  Ameri- 
can lists. 

The  species  in  the  genus  are  superficially  so  similar  in  color  and 
general  characteristics  that  there  has  been  much  confusion  in 
naming  them.  The  descriptions  have  been  made  in  several  cases 
from  a  single  specimen,  without,  therefore,  a  consideration  of  varia- 
tions within  the  species.  They  have  dealt  with  comparisons  that 
can  be  appreciated  only  after  long  familiarity  with  the  insects,  and 
on  this  account  many  errors  of  determination  have  arisen.  The 
most  unfortunate  circumstances  attending  the  study  of  North 
American  Ranatra  has  been  the  failure  to  recognize  the  identity  of 
Ranatra  jusca  P.  B.  1805  and  Ranatra  nigra  H.  S.  1853,  the  de- 
scriptions of  which  are  inadequate,  and  therefore,  since  the  types  are 
not  available,  can  be  interpreted  authoritatively  only  through  the 
examination  of  abundant  and  representative  material  from  the 
entire  country. 

After  studying  long  series  of  specimens  representing  a  wide  dis- 
tribution, the  writer  has  been  compelled  by  the  evidence  before  him 
to  place  a  new  interpretation  upon  our  North  American  Ranatra. 
This  lie  has  been  reluctant  to  do,  because  it  involves  the  renaming 
of  our  commonest  two  species.  Stability,  however,  can  never  be 
attained  in  this  group  by  postponing  or  ignoring  the  evidence  that 
eventually  must  prevail. 

Ranatra  jusca  P.  B.  was  described  and  figured  in  color,  natural 
size,  by  Palisot  de  Beauvois,  in  his  'Tnsectes  recueillis  en  Afrique  et 
in  Amerique,  dans  les  Royaumes  d'6ware  et  de  Benin,a  Saint  Domin- 
que  et  dans  les  Etats  Unis,  pendant  les  annees  1786-1797." 

The  figure  is  not  amplified  nor  enlarged  as  suggested  by  Doctor 
Montandon  (Bulletin  Soc.  Sci.  Bucharest,  vol.  XIX,  1910),  who  was 
in  error  in  endeavoring  to  make  R.  kirkaldyi  Bueno,  our  smallest 
North  American  Ranatra,  fit  a  drawing  that  has  the  dimensions  of 
our  largest  species.  He  was  mistaken  in  believing  R.  kirkaldyi 
Bueno  a  synonym  of  R.  jusca  P.  B.,  for  there  are  two  convincing 
reasons  for  believing  R.  jusca  P.  B.  is  represented  natural  size. 

First.  Palisot  de  Beauvois  in  his  "Discours  Preliminaire,"  page 
XV,  says,  "J'ai  adopte,  pour  la  grandeKur  des  figures,  un  plan  uni- 
forme,  et  qui  m'a  semble'plus  commode,  c'est-a-dire,celle  d'un  pouce 
pour  tous  les  Insectes  plu^  petits  que  cette  mesure  adoptee,  en  plac- 


Hungerford:    Nepid.k  in  America.  435 

ant  a  cote  une  ligne  perpendiculaire- de  la  grandeur  reerdc  Tanimal. 
Ceux  (lui  excedent  un  ponce  seront  figures  de  Icur  grandeur  na- 
turelle." 

Second.  The  plate  which  bears  the  figure  of  Ranatra  fusca  has 
upon  it  three  figures  natural  size  (one  of  which  is  Ranatra  fusca)' 
and  two  figures  enlarged.  These  two  have  the  true  length  indicated 
by  a  line,  and  one  of  these,  Naucoris  jemorata,  of  w^iicli  the  writer 
has  specimens,  fits  the  line  which  was  drawn  to  indicate  the  true 
length.  There  are  on  the  plate  three  figures  of  Belostoma  which 
Palisot  calls  "Nepa,"  male  and  female,  of  "Nepa  suhspinosa,"  nat- 
ural size,  which  are  not  accompanied  by  a  "line  of  true  length,"  and 
Neya  minor,  accompanied  by  a  line.  The  drawing  of  Neya  minor  is 
as  large  as  those  of  Nepa  subspinosa,  and  yet  the  author  distinctly 
says  that  Nepa  minor  is  smaller.  The  line  indicates  its  true  length, 
and  the  inevitable  conclusion  is  that  the  figure  of  Ranatra  fusca  is 
natural  size.  Indeed,  a  comparison  of  drawings  with  the  specimens 
throughout  the  book  shows  the  author  consistently  followed  his 
stated  policy  of  drawing  large  insects  natural  size  and  indicating 
enlargements  by  lines,  whenever  made. 

Palisot's  insects  were  taken  between  the  years  1786  and  1797. 
Those  from  the  United  States,  then,  were  taken  from  somewhere  in 
the  eastern  United  States,  for  the  territories  of  the  United  States  at 
that  time  were  bounded  on  the  north  by  Canada,  on  the  west  by  the 
Alississippi  river,  and  on  the  south  by  the  north  line  of  Florida. 
Ranatra  fusca,  therefore  was  collected  within  these  limits. 

Palisot,  in  his  "Discours  Pr^liminaire,"  alludes  to  an  account  of  his 
travels  as  being  in  press.  I  have  not  located  this,  but  have  secured 
two  brief  biographical  sketches  of  him;  one  from  "La  Grande  Ency- 
clopedie,"  vol.  XXV,  and  the  other  more  complete,  from  P.  Larousse, 
"Dictionnaire  du  XIX  Seicle  12,"  page  66.  Neither  of  these  mention 
a  book  of  his  travels.  From  these  accounts  we  learn  that  he  came  to 
Philadelphia  first  in  1791  to  secure  assistance  against  a  Negro  up- 
rising in  San  Domingo,  where  he  was  taking  part  in  govermental 
affairs.  He  seems  to  have  been  in  the  United  States  for  some  time, 
for  upon  his  return  to  San  Domingo  he  narrowly  escaped  death  at 
the  hands  of  the  blacks  (17931.  He  escaped  to  Philadelphia  with- 
out funds  and  supported  himself  there  by  giving  lessons  in  music 
and  language.  Finally  he  secured  help  from  France  to  make  a 
scientific  voyage  in  North  America  for  new  collections,  and  returned 
to  his  own  country  in  1798.  He  died  in  1820.  These  brief  facts 
indicate  that  most  of  his  days  in  the  United  States  were  spent  about 
Philadelphia,  and  it  seems  to  me  probable  that  many  of  his  insects 


436  The  University  Science  Bulletin. 

were  collected  in  that  region.     The  possibility  of  his  R.  fusca  not 
being  known  to  us  in  nature  seems  to  me  quite  remote. 

According  to  the  figure,  Ranatra  fusca  of  Palisot  de  Beauvois  is 
a  large,  robust  insect  with  broad  anterior  femora  and  short  legs  and 
respiratory  tube.  The  cjuestion  is,  what  insect  of  the  range  indi- 
cated has  these  characters?  After  careful  consideration  of  the 
problem  I  must  conclude  that  the  only  one  with  the  proper  size, 
robustness,  broad  femora  and  short  legs  is  Ranatra  americana 
Montd.  It  has  the  variable  respiratory  tube,  usually  longer  than 
Palisot's  figure,  but  not  uncommonly  as  short,  and  in  every  case 
shorter  than  the  body.  The  failure  of  the  artist  to  indicate  any 
apical  tooth  on  the  anterior  femur  is  readily  understood,  since  we 
know  that  this  character  is  often  obscured  by  a  marginal  fringe  of 
pile  so  completely  as  to  escape  any  but  the  closest  scrutiny.  It  is 
indeed  a  somewhat  variable  structure;  in  some  examples  much  re- 
duced, and  in  others  plainly  visible.  The  writer  has  in  his  collection 
specimens  which  fit  Palisot's  figure  almost  exactly. 

The  second  species  with  which  we  are  concerned  is  Ranatra  nigra 
H.  S.  This  species  was  described  in  1853  from  America  by  Herrich 
Schaffer  in  his  "Die  Wanzenartigen  Insecten."  It  was  described  as 
being  from  2  to  21/8  inches  long  from  beak  to  tip  of  respiratory 
tubes,  with  the  respiratory  tubes  not  much  over  half  as  long  as  the 
body;  indeed,  he  says  they  were  three-fourths  of  an  inch!  Now  the 
only  species  we  have  which  in  a  series  of  specimens  has  a  respiratory 
tube  averaging  three-fourths  of  an  inch  long  is  R.  protensa  Montd. 
Doctor  Montandon  described  his  R.  protensa  from  a  single  large 
female,  which,  because  of  its  very  yellowish  color  and  shorter  limbs, 
did  not  suggest  R.  nigra  to  his  mind.  Indeed,  considered  alone,  it  is 
not  at  all  to  be  expected  that  it  would.  R.  nigra  was  described  as 
having  the  structure  of  thorax  and  relative  length  of  limbs  as  in  i?. 
elongata.  Now,  R.  elongata  has  very  elongate  hind  femora,  the  tips 
surpassing  the  last  abdominal  suture  by  a  considerable  distance. 
The  length  of  limb  in  R.  protensa  Montd.  is,  on  the  whole,  not  as 
great  as  in  R.  elongata,  but  is  strikingly  longer  than  in  R.  linearis 
L.,  with  which  he  compared  it  in  size,  and  this  suggested  R.  elongata. 
Indeed  in  many  specimens  of  R.  protensa  Montd.  the  hind  femora 
surpass  the  last  abdominal  suture.  The  front  legs  of  R.  protensa 
are  long  and  very  slender  and  the  thorax  is  more  like  that  of  R. 
elongata  than  of  R.  linearis.  Doctor  Montandon  said  that  he  had 
never  seen  any  specimens  from  North  America  with  legs  as  in 
elongata.    We  have  three  species  with  limbs  relatively  as  long. 


Huxcerford:    Nepid.i-:  ix  America.  437 

It  seems  scarcely  necessary  to  mention  that  the  color  implied  by 
the  name  7iigra  has  nothing  to  do  with  the  case,  for  black  coloration 
occurs  in  all  the  species,  due  to  one  of  two  causes — either  dark  in- 
crustations or  deposits  upon  the  integument,  or  dark  discoloration 
due  to  failure  in  drying  out  the  specimens.  Either  of  these  causes 
might  account  for  a  uniform  dark  or  black  color  in  a  given  series 
of  insects. 

There  are  other  facts  relative  to  the  identity  of  these  species 
which  will  be  presented  under  notes  after  the  various  descriptions, 
to  which  the  I'eader  is  referred  for  further  evidence.  Aside  from  the 
points  mentioned,  the  probability  of  our  two  commonest  and  wide- 
spread species  being  first  described  is  of  itself  very  great. 

The  list  of  the  species  of  Ranatra  now  known  from  America  north 
of  Mexico  is  as  follows: 

Ranatra  fusca  P.  B.  1805  =  Ranatra  americana  Montd.  1910. 

Ranatra  nigra  H.  S.  185^  ^  Ranatra  protensa  Montd.  1910. 

Ranatra  quadridentata  Stal  1861. 

Ranatra  kirkaldyi  Bueno  1905  and  its  variety  hofjmanni,  n.  var. 

Ranatra  brevicollis  Montd.  1910. 

Ranatra  buenoi,  sp.  new  =  Ranatra  fusca  Bueno  and  Mondt.  in 
part. 

Ranatra  drakei,  sp.  new. 

Ranatra  australis,  sp.  new. 

It  will  be  noted  that  R.  quadridentata  Stal,  completely  submerged 
in  Van  Duzee's  catalogue  under  R.  americana  Montd.,  is  restored. 
Dr.  F.  H.  Snow,  in  Trans.  Kan.  Acad.  Sci.,  vol.  XX,  pt.  1,  p.  153, 
1906,  w^as  not  writing  about  the  species  which  Doctor  Montandon 
later  described  as  R.  americana,  but  another  which  we  believe  to  be 
R.  quadridentata  Stal,  and  should  not  have  been  synonymized  by 
Van  Duzee.  It  will  also  be  seen  that  R.  annulipes  Stal  is  omitted. 
This  R.  annulipes  Stal  is  a  very  distinct  species  and  not  to  be  mis- 
taken for  any  the  writer  knows  from  our  range.  Doctor  Montandon 
has  established,  through  the  examination  of  types  and  type  ma- 
terial, that  R.  fabricii  Guer.  is  the  same  as  R.  annulipes  Stal.  Fig- 
ures of  this  species  are  given  in  this  paper,  because  it  has  been  cited 
as  coming  from  our  range. 

The  variety,  edentata  Montd.,  of  R.  americana  Montd.,  is  as- 
sumed to  be  an  americana  with  attenuated  apical  tooth  on  front 
femur.    Whether  it  should  be  recognized  as  a  variety  is  questionable. 

The  entire  question  of  the  correct  names  for  the  Ranatra  was,  as 
the  writer  has  stated,  opened  by  the  necessity  of  naming  four  species 


438  The  University  Science  Bulletin. 

from  Kansas.  The  problem  was  undertaken  with  no  preconceived 
notions  in  the  matter,  except,  indeed,  a  leaning  toward  accepting 
Doctor  Montandon's  studies,  since  he  is  a  great  scholar  in  the 
Cryptocerata  and  was  followed  by  Van  Duzee  in  his  catalogue.  The 
results  herewith  presented  may  be  received  with  some  irritation, 
because  of  the  revolution  necessary  in  nomenclature,  but  it  seems 
best  for  us  to  get  as  near  the  truth  as  the  evidence  permits. 

In  an  endeavor  to  fix  the  limits  of  the  species,  the  writer  has 
studied  the  characters  used  by  the  best  systematists  in  the  group 
and  in  addition,  antennal  and  genital  structures.  The  relative  length 
of  body  parts,  the  comparative  size  of  the  eyes,  the  relative  length 
of  the  limbs  and  of  the  caudal  filaments,  the  shape  of  the  sternum 
and  the  form  of  the  front  femora  have  been  in  general  use  in  the  at- 
tempts to  define  nature's  species  in  this  genus.  Species  are  expected 
to  vary  within  limits,  and  breeding  experiments  with  insects  show 
them  to  vary,  not  only  in  single  characters,  but  in  linked  characters, 
and  in  order  to  steer  a  fair  course  between  ''lumping"  and  ''split- 
ting" species,  it  is  much  better  to  use  a  combination  of  three  or  more 
characters  than  to  depend  too  much  upon  any  single  one.  The  use, 
therefore,  of  the  characters  of  the  antennae  and  genitalia  is  a  ma- 
terial aid  in  the  classification  of  this  difficult  group. 

NOTES  ON  TERMS  USED  IN  KEYS  AND  DESCRIPTIONS. 

The  apical  tooth  of  the  front  femur  is  on  the  inside  edge  of  the 
front  femur  near  the  attachment  of  the  tibia  (see  fig.  2-A,  pi. 
XLIV).  The  antennae  are  hidden  in  pockets  beneath  the  eyes  (see 
fig.  9^  pi.  XLV).  The  prothorax  of  Ranatra  is  divided  into  two 
parts,  anterior  and  posterior,  by  transverse  lateral  grooves  (see  fig. 
2-G,  pi.  XLIV).  The  tylus  and  jugae  are  figured  on  plate  XLV, 
figure  12.  The  metaxyphus  extends  back  between  the  hind  coxae 
as  a  part  of  the  metasternum.  The  last  abdominal  segment  is  desig- 
nated as  the  "genital  segment,"  and  its  ventral  plate  in  both  sexes  as 
the  "operculum." 

In  this  paper  I  have  not  distinguished  between  the  genital  seg- 
ment and  the  last  abdominal  segment,  as  is  often  done  with  some 
other  Heteroptera,  where  the  last  abdominal  segment  is  considered, 
the  segment  just  in  front  of  the  genital  segment.    I  refer  to  this  as 
the  penultimate  abdominal  segment  (see  figs.  10  and  11,  pi.  XLV). 

The  male  genital  capsule  of  which  I  speak  is,  I  take  it,  homolo- 
gous with  what  G.  C.  Crampton  in  the  Bulletin  of  the  Brooklyn 
Entomological  Society,  volume  XVII,  pages  45-55,  calls  the  hypan- 
drium,"  and  the  claspers  correspond  to  his  "styli,"  or  "gonostyles." 


Hungerford:    Nepid.e  in  America.  439 

TECHNIQUE  USED. 

The  antennee  arc  often  covered  with  debris  that  should  be  re- 
moved. If  the  antennae  are  still  difficult  to  see,  the  specimen  may  be 
relaxed  and  antennae  moved  into  view  with  dissecting  needle.  The 
genital  capsule  of  the  male  lies  in  the  last  abdominal  segment,  en- 
tirely hidden.  To  examine  this,  relax  the  specimen  in  a  moist  cham- 
ber, then  holding  the  specimen  on  its  right  side  in  the  left  hand, 
lower  the  operculum  with  a  dissecting  needle  and  draw  out  the 
capsule.  A  needle  with  just  the  very  tip  turned  at  right  angle  is 
best  to  use.  It  can  be  inserted  alongside  the  capsule,  turned  so 
that  the  hook  is  against  the  capsule,  and  slowly  withdrawn,  bringing 
with  it  the  desired  organ.  If  the  respiratory  filaments  start  to  come 
also,  loosen  the  capsule  from  them  by  inserting  the  needle  between 
capsule  and  filaments  on  each  side.  The  capsule  may  be  removed 
entirely  and  mounted  upon  a  card  below  the  insect,  or  simply  drawn 
into  view.  The  insects  are  not  in  the  least  damaged,  and  the  male 
capsules  may  be  taken  out  and  operculum  pressed  back  into  place 
by  a  careful  person  without  it  being  detected  by  the  closest  exam- 
ination of  the  exterior. 

KEY   TO  THE  SPECIES. 

A.  Antennae  simple,  distal  end  of  the  penultimate  segment  without  lateral 
prolongation.  Front  femora  broad  and  stout  and  not  narrowed  near 
middle. 

B.    Front  femora  without  apical  tooth  or  marked  sinuosity. 

R.  kirkaldyi  Bueno. 
BB.    Front  femora  with  apical  tooth  or  a  marked  sinuosity. 

R.  kirkaldyi  Bueno. 
var.  hoffmanni,  new. 

AA.   Antennae  with  distal  end   of  penultimate  segment  with  a   lateral  pro- 
longation.   Front  femora  somewhat  narrowed  in  their  middle  section. 

B.  The  lateral  prolongation  of  the  penultimate  segment  of-  antennae 
not  greater  than  one-half  of  the  length  of  the  ultimate  segment. 
Front  femora  very  slender  and  without  apical  tooth. 

C.   Prosternum  with  a  single  wide,  deep  longitudinal  trough.    Eyes 
veiy  prominent.  .  R.  buenoi,  sp.  new. 

{=R.  jusca  Bueno.) 
CC.   Prosternum    without    the    deep    trough,    but    possessing    two 
longitudinal  depressed  lines  characteristic  of  most  species  of 
Ranatra.  R.  nigra  H.  S. 

{=R.  protensa  Montd.) 
BB.   The  lateral  prolongation  of  the  penultimate  segment  of  antennae 
greater  than  one-half  the  length  of  the  ultimate  segment. 
C.    Sides  of  body  (connexivum)  embracing  the  operculum  of  the 
genital  segment  at  its  extremity  (see  fig.  11,  pi.  XLV), 

R.  annulipes  Stal. 
{=:R.  fabricii  Gu^r.) 


440  The  University  Science  Bulletin. 

CC.  Sides  of  body  not  embracing  the  genital  operculum  at  its  ex- 
tremity (see  fig.  10,  pi.  XLV). 
D.  Pronotum  broad  and  short,  anterior  enlargement  sub- 
equal  in  width  to  entire  head.  Anterior  femur  broad  and 
stout  without  apical  tooth.  Median  tooth  half  way  be- 
tween trochanter  and  tibial  joint  (measured  on  inner 
edge).  Metaxyphus  very  long,  extending  beyond  middle 
of  hind  coxae,  nearly  attaining  abdomen. 

R.  brcvicoUis  Montd. 
DD.   Pronotum  more  slender.     The  median  tooth  of  anterior 
femur   nearer   distal    than   proximal    end.      Metaxyphus 
not  often  extending  bej'ond  the  middle  of  hind  coxae. 

E.    Front  femur  broad  without  apical  tooth. 

F.    Jugae  of  head  more  prominent  than  tylus. 

R.  mistralis,  sp.  new. 
FF.   Tylus  fully  as  prominent  as  jugae. 

R.  jusca  P.  B. 
(var.  edentula  Montd.) 
EE.    Front  femur  with  apical  tooth. 

F.  Anterior  portion  of  prothorax  fully  twice  as 
long  as  thickened  posterior  portion.  Eyes 
large,  plainly  greater  than  interocular  space. 
Jugae  of  head  prominent.  Front  femur  long 
and  slender.  Hind  femur  surpassing  middle  of 
penultimate  abdominal  segment.  Caudal  fila- 
ments as  long  as  body.  R.  drakei,  sp.  new. 
FF.  Anterior  portion  of  prothorax  shorter  than 
above.  Eyes  not  much,  if  any,  greater  than 
interocular  space.  Jugae  not  so  prominent. 
Front  femur  fairly  stout.  Middle  and  hind 
femora  short,  not  attaining  middle  of  penulti- 
mate segment. 

G.  Eyes  prominent,  plainly  transverse,  fully 
as  large  as  interocular  space.  Anterior 
part  of  pronotimi  somewhat  flattened 
dorsoventrallv.     Hind  femora  short. 

R.  jusca  P.  B. 
(=:R.  americana  Montd.) 
GG.   Eyes  not   prominent,  neither  transverse 
nor  as  large  as  interocular  space.     An- 
terior part  of  pronotum  more  cylindrical. 
R.  quadridentata  Stal. 

.  Ranatra  kirkaldyi  Bueno. 

Bueno,  J.  R.  de  la  Torre,  Can.  Ent.  XXXVII,  p.  187,  1905. 

(= -R.  fusca  Montd.,  Bui.  Soc.  Sci.  Bucharest,  XIX,  p.  3;   1910.) 

Original  description: 

Abdominis  dorsum  orange  brown;  eyes  small,  not  very  prominent;  pro- 
thorax much  constricted  at  the  middle,  bisulcate  beneath;  wings  smoky; 
anterior  femora  broad  with  a  prominent  tooth  near  the  middle,  otherwise 
smooth;  posterior  tarsi  extending  beyond  the  middle  of  the  air  tube;  air 
tube  shorter  than  the  length  of  the  abdomen;  legs  banded.  Length  from  tip 
of  abdomen  to  tip  of  rostrum,  male,  23  mm.  to  26.4  mm.;  female,  27  to  31  mm. 


Hungerford:    Nepid.e  in  America.  441 

Notes.  Tliis  is  the  species  that  Doctor  Montandon  has  con- 
sidered a  synonym  of  I\\  fusca  P.  B.*  Its  much  smaller  size  (7?. 
fusca  P.  B.  measures  37  mm.)  is  not  the  only  reason  for  believing 
otherwise.  Tlie  anterior  femora  of  R.  kirkaldyi  Bueno  are  not  con- 
stricted near  the  middle  tootli  as  they  are  in  R.  fusca  P.  B.  The 
front  legs  and  respiratory  tube  are  both  shorter  relatively  than  in 
Palisot's  figure.  This  is  the  smallest  known  species  in  North  Amer- 
ica. It  is  a  characteristic  species,  quite  unrelated  and  distinct  from 
the  others.  In  the  first  place,  the  antennae  are  much  reduced.  They 
are  straight,  lacking  any  projections  from  the  penultimate  segment, 
and  often  lack  anything  more  than  a  constriction  to  distinguish  the 
terminal  segment.  The  ultimate  segment  is  sometimes  fused  with 
the  penultimate  (see  figs.  9  and  11,  pi.  XLVII),  but  all  stages  of 
separation  can  be  found.  The  male  genital  claspers  are  also  very 
distinct,  as  shown  by  figure  5  on  plate  XLVI. 

Specimens  of  this  species  taken  by  Beamer  and  Hungerford  in 
Cherokee  county,  Kansas,  were  identified  by  Mr.  J.  R.  de  la  Torre 
Bueno.  They  agree  with  the  original  description  in  lacking  any 
apical  tooth  or  sinuosity  near  the  apex  of  the  femora.  They  measure 
from  tip  of  beak  to  base  of  filaments,  from  24  to  30  mm.,  wdth  the 
filaments  from  14  to  17  mm.  long.  I  have  seen  the  true  R.  kirkaldyi 
Bueno  from  New  York. 

In  addition  to  the  above  Kansas  series,  I  have  before  me  the 
following  series  of  127  specimens  as  follows: 

8  Rockbridge,  Ohio,  9-30-16,  C.  J.  Drake. 

1  Rocky  Mount,  N.  C,  10-19-16,  R.  W.  Leiby. 

3  St.  Paul,  Minn.,  Phalen  Lake,  6-19-21,  H.  B.  Hungerford. 

1  St.  Paul,  Minn.,  Lake  Johanna,  6-26-21,  H.  B.  Hungerford. 

4  Ramsey  county,  Minnesota,  1-9  22,  W.  E.  Hoffman. 

7  St.  Paul,  Minn.,  Lake  Johanna,  10-13-22,  W.  E.  Hoffman. 

9  St.  Paul,  Minn.,  Lake  Johanna,  10-14-22,  W.  E.  Hoffman. 
32  St.  Paul,  Minn.,  Lake  Johanna,  W.  E.  Hoffman. 

3  Lincoln,  Neb.,  1  male,  2  females,  W.  E.  Hoft'man. 

39  St.  Paul,  Minn.,  Water  Supply  Canal,  10-18-21,  W.  E.  Hoffman. 
16  St.  Paul,  Minn.,  Lake  Johanna,  9-24-21,  W.  E.  Hoffman. 

4  St.  Paul,  Minn.,  Lake  Johanna,  9-17-21,  W.  E.  Hoffman. 

I  have  also  seen  a  series  of  170  taken  by  Doctor  Drake  in  Ohio. 

These  Minnesota  specimens  have  certain  characteristics  which 
set  them  clearly  apart  from  the  Kansas  series,  yet  there  can  be  no 
mistake  in  considering  them  as  belonging  to  the  same  species.  Since 
I  have  had  a  fairly  good  series  of  these  insects,  which  are  smaller 

*See  plate  XLIX,  figures  1  and  2. 


442  The  University  Science  Bulletin. 

than  the  Kansas  forms,  more  robust,  with  front  femur  relatively- 
thicker  and  possessing  a  more  or  less  well-marked  sinuosity  near  the 
inner  apex,  with  antennae  which,  on  the  whole,  are  blunter  at  tip, 
and  the  subapical  tooth  of  the  claspers  of  male  slightly  more  slen- 
der, it  seems  wise  to  consider  them  a  variety,  for  which  I  propose 
the  name  Ranatra  kirkaldyi  var.  hoffmanni.  It  will  be  noted  that 
Mr.  W.  E.  Hoffman  has  provided  me  with  the  longest  series  of  this 
variety.  In  a  series  as  large  as  this,  it  would  seem  that  if  the  variety 
were  not  fixed  there  should  be  found  forms  like  the  true  kirkaldyi, 
which  is  described  as  having  "anterior  femora  broad  with  a  promi- 
nent tooth  near  the  middle,  otherwise  smooth." 

Ranatra  buenoi,  sp.  new. 

=  R.  fusca  Bueno,  Can.  Ent.  XXXVII,  p.   188,  1905. 

Bueno  and  Brimley,  Ent.  News,  XVIII,  p.   438,  1907. 

Bueno,  Ent.  News,  XXXII,  p.  273,  1921. 

Montandon ;   determination  in  U.  S.  National  Museum. 

Size.  Length  from  tip  of  beak  to  tip  of  abdomen,  32  mm.  to  38  mm.;  caudal 
filaments,  22  mm.  to  27  ami.  long.. 

Color.  From  light  to  very  dark  fuscous;  top  of  abdomen  orange  and  black; 
middle  and  hind  legs  of  lighter  forms  banded. 

Shape.  Long  and  slender,  prothorax  long  and  slender;  eyes  very  prominent 
and  large;  limbs  all  very  slender  and  very  long;  the  under  side  of  prothorax 
with  a  single  broad  and  deep  sulcus,  which  distinguishes  this  species  from  any 
of  our  other  forms  (see  fig.  6,  pi.  XLVIII). 

Structural  peculiarities.  The  eyes  very  prominent,  transverse  diameter 
greater  than  the  interocular  space;  tjdus  longer  than  jugae  and  as  prominent 
as  these  parts,  which  are  of  medium  development;  antennae  with  the  lateral 
prolongation  of  penultimate  segment  not  more  than  half  the  length  of  ulti- 
mate; prothorax  slender,  the  anterior  portion  measured  on  the  median  dorsal 
line,  twice  the  length  of  the  posterior  swollen  part  (the  well-marked  long  trans- 
verse lateral  grooves  used  as  di\ading  line) ;  scutellum  somewhat  elevated  and 
slender  and  the  length  of  the  abdomen  is  to  pronotum  as  2^  is  to  1;  respira- 
tory filaments  surpassing  front  margin  of  scutellum  when  brought  forward; 
claspers  of  male  genital  bulb  very  distinct;  subapical  tooth  greatly  reduced 
(see  fig.  1,  pi.  XLVI) ;  all  the  limbs  strikingly  long  and  slender,  especially  the 
anterior  femora  (see  fig.  1,  pi.  XLVIII) ;  no  apical  tooth  on  anterior  femur,  the 
other  tooth  much  nearer  the  apex  than  the  base,  the  basal  part  being  at  least 
IV2  times  as  long  as  the  part  lying  before  the  tooth;  the  coxa  two-thirds  as 
long  as  the  femur. 

The  middle  and  hind  legs  long,  the  distal  ends  of  hind  femora  surpassing 
the  last  abdominal  suture  and  often  nearly  attaining  caudal  end  of  genital 
segment,  while  the  middle  femora  frequently  attain  or  surpass  the  caudal 
margin  of  penultimate  abdominal  segment;  the  distal  ends  of  middle  and 
hind  legs  almost  attaining  tip  of  respiratory  filaments;  the  relative  lengths 
of  femora  to  tibiae  are  about  as  16  is  to  19  for  the  middle  leg  and  as  16  is  to 
22  for  hind  leg.  The  tarsi  are  small  relatively,  a  little  less  than  one-sixth  as 
long  as  their  tibiae. 


Hungerford:    Nepid.k  in  America.  443 

Notes.    Described  from  the  following  series: 

Male  holotype,  Colorado  county,  Texas,  June  24,  1922;   Mrs.  Grace  Wiley, 

collector. 
Female  allotype,  Colorado  county,  Texas,  June  24,  1922;  Mrs.  Grace  Wiley, 

collector. 
Paratypes  as  follows: 

2  9  $    Raleigh,  N.  C,  July  10,  1902;  F.  Sherman,  jr.,  collector. 

$    Mound,  La.,  November  6,  1918. 
9    Aberdeen,  Miss.;  Dr.  Carl  Drake. 

3  9  9   and  IS   Leiand,  Miss.,  September  16,  1921;  C.  J.  Drake. 

9   Creve  Coeur  Lake,  Mo.,  May  15,  1911;  J.  F.  Abbott. 
$   Dime  Box,  Tex.,  July  20,  1911;  C.  T.  Atkinson,  collector. 
17   ,J  5   and  22   9  9   Gainesville,  Fla.,  June  19,  1918;  Carl  Drake. 

Holotype  and  allotype  in  University  of  Kansas  museum;  para- 
types in  collection  of  Dr.  Carl  Drake,  J.  R.  de  la  Torre  Bueno,  and 
the  author. 

This  species  has  been  named  for  Mr.  J.  R.  de  la  Torre  Bueno, 
who  pointed  out  its  structural  characters  in  1905.*  It  has  also 
been  named  by  some  workers  R.  fusca  P.  B.,  and  by  at  least  one 
R.  nigra  H.  S.  It  is  clearly  impossible  for  it  to  be  either.  The  error 
of  considering  it  R.  fusca  P.  B.  has  been  due  to  the  inadequate  de- 
scription of  R.  fusca  P.  B.  and  to  the  fact  that  the  original  text  with 
illustration  has  not  been  accessible  to  many,  if  any,  American  stu- 
dents of  this  group.  By  taking  Palisot  de  Beauvois'  figure  of  his 
R.  fusca  and  comparing  it  with  the  species  above  described,  it  will 
be  seen  at  once  that  the  species  are  not  the  same.  The  long,  slender 
limbs,  the  great  eyes,  the  long  thorax,  the  relative  position  of  the. 
tooth  on  the  front  femur,  and  the  relation  in  length  between  femora 
and  tibiae  of  the  legs,  preclude  the  possibility  that  Palisot's  artist 
had  R.  buenoi,  sp.  new,  before  him.  The  front  legs  of  R.  fusca  P.  B. 
are  stout,  not  exceedingly  slender.  The  median  tooth  of  front  femur 
is  nearer  the  middle  than  in  R.  buenoi,  sp.  new.  The  anterior  part 
of  prothorax  is  less  than  twice  the  swollen  part.  The  middle  femora 
and  tibise  are  nearly  the  same  length  as  they  are  in  R.  americana 
Montd.,  for  instance,  and  not  considerably  different  in  length  as  in 
R.  buenoi.  The  hind  femora  are  considerably  shorter  relative  to  the 
abdomen  in  R.  fusca  P.  B.  The  tarsus  of  R.  fusca  P.  B.  is  larger, 
being  one-fifth  as  long  as  its  tibia  and  not  one-sixth  as  in  R.  buenoi 
sp.  new.  In  other  words,  its  proportions  fit  another  American  spe- 
cies, but  not  R.  buenoi. 

This  species  is  not  R.  nigra  H.  S.,  for  R.  nigra  H.  S.  has  uni- 
formly short  respiratory  tube  of  three-fourths  inch  and  measures 

*See  Can.  Ento.,  vol.  XXXVII,  p.  188;    1905. 


444  The  University  Science  Bulletin. 

over  all  from  tip  of  beak  to  end  of  filaments  2  to  2%  inches,  while 
this  species  has  a  tube  averaging  a  full  inch  and  varies  in  size  from 
2^8  to  21/2  inches.  The  shape  of  the  eyes  as  shown  in  Herrich- 
Schaffer's  figure,  is  very  different,  and  the  size  which  he  says  is 
"not  larger  than  R.  linearis,"  would  at  once  eliminate  it,  for  in 
R.  buenai  they  are  very  large,  but  not  so  large  as  in  R.  elongata, 
specimens  of  which  I  have  seen. 

For  illustration  of  this  species,  see  figures  1  and  2,  plate  XLVI; 
fig.  12,  plate  XLVII;  figure  1,  plate  XLVIII;  and  figure  5,  plate 
XLIX.  This  species,  as  Bueno  has  written,  is  southern  in  distribu- 
tion. Besides  the  localities  above  cited,  I  have  seen  it  from  Way- 
cross,  Ga.,  taken  by  J.  C.  Bradley,  U.  S.  N.  M. 

Ranatra  nigra  Herrich-Schaffer. 

Henich-Schaffer,  G.  A.  W.,  Wanzenartigen  Insecten,  IX,  p.  32  ;    1853. 
(^  Runatra  proie»sa  Montd.   and  all  subsequent  writers.) 

Original  description : 

R.  nigra  m.  tab.  290,  fig.  L,  2  =  2%".  Nigra,  tubis  respiratoribus  longi- 
tiidinem  dimidiam  corporis  parum  superantibus.  Grosse  fast  von  R.  linearis, 
schwarzer,  die  Aiigen  night  grosser  aber  seitlich  mehr  vortretend,  der  Scheitel 
daher  breiter.     Ban  des  Thorax   imd   Liingenverhaltnisse   der  Beine  wie  bei 

R.  elongata.    Aus  Anierika. 

In  addition  to  the  above,  and  in  the  discussion  relative  to  the 
genus  Ranatra  and  the  species  R.  elongata,  R.  filiformis,  R.  linearis, 
macrophthalma  and  R.  nigra,  he  gives  notes  of  comparison  which 
throw  further  light  upon  his  species.  He  says  R.  elongata  has  the 
longest  respiratory  tube  (134  inches)  and  R.  nigra  has  the  shortest 
{%  inch).  The  color  is  darkest  in  R.  nigra,  "die  Augen  sind  am 
grossten  bei  macrophthalma,  am  kleinsten  bei  linearis,  am  rundes- 
ten  bei  letzerer,  am  meisteiijaie  Quere  gezogen  bei  nigra."  His  state- 
ment of  length,  2  to  2^8  inches  from  tip  of  beak  to  tip  of  tube  with 
"Athmungsrohren  nicht  viel  iiber  halb  so  lang  als  der  Korper,"  in- 
dicate that  he  had  a  series  of  the  insects. 

Notes.  The  only  species  of  our  country  which  agrees  in  size  and 
uniformly  short  length  of  respiratory  tube  with  the  description  of 
R.  nigra  is  the  insect  we  know  under  the  name  R.  protensa  Montd. 
Doctor  Montandon  had  before  him  a  single  large,  fairly  short- 
limbed  female,  which  is  his  type  deposited  in  the  United  States  Na- 
tional Museum.  This  $  was  collected  by  Wm.  H.  Ashmead  on 
Long  Island.  The  respiratory  tube  is  shorter  than  the  abdomen. 
The  legs  extended  would  surely  reach  the  tip  of  the  respiratorv  tube. 

Ranatra  nigra  is  quite  common  over  our  country  everywhere  I 
have  collected.     It  is  most  unfortunate  that  it  was  described  from 


Hungerford:    Nepid.e  in  America.  445 

blackened  specimens,  for  indeed  it  is  our  lightest-colored  species, 
being  of  a  yellowish  color  with  a  trace  of  green  throughout.  This 
species  has  very  prominent  mesocoxae  and  metacoxse,  which  are 
angular,  having  a  slight  tubercle  on  the  inner  side.  The  breast,  or 
prothorax  in  front  of  the  mesocoxal  elevations,  is  constricted  and 
small,  making  the  coxal  elevations  very  pronounced. 

The  United  States  National  Museum  possesses,  besides  the  type, 
specimens  from  Virginia,  West  Virginia,  Pennsylvania,  Maryland, 
Arkansas  and  Florida. 

I  have  before  me  a  series  of  seventy-nine  specimens  from  Doni- 
phan county,  Kansas,  taken  by  Robert  Guntert  and  W.  J.  Brown; 
ten  from  Douglas  county,  Kansas,  and  forty  specimens  taken  by 
Mr.  W.  E.  Hoffmann  and  myself  from  the  following  places  in 
Minnesota:  Lake  Johanna,  Phalen  lake  and  Minnehaha  creek,  all 
near  the  twin  cities.  We  have  also  taken  it  in  other  parts  of  the 
state.  Careful  study  of  ample  material,  well  distributed,  shows 
that  by  far  the  commonest  size  is,  total  length,  two  inches,  with  re- 
spiratory tube  three-fourths  inch!  The  length  of  hind  femur  rel- 
ative to  abdomen  varies,  sometimes  attaining  the  front  margin  of 
the  genital  segment  and  sometimes  falling  somewhat  short  of  this. 
The  eyes  are  transverse  and  somewhat  protuberant.  The  jugse  fit 
closely  against  the  tylus.  The  lateral  prolongation  of  the  penulti- 
mate segment  of  the  antennae  is  less  than  half  the  ultimate.  The 
front  femora  are  very  slender  and  coxae  long.  On  page  156  of  the 
University  of  Kansas  Science  Bulletin,  vol.  XI,  where  I  give  an 
English  translation  of  Doctor  Montandon's  description  in  French 
of  R.  protensa,  I  should  have  written,  "Anterior  femora  quite  slen- 
der, but  scarcely  a  fifth  longer  than  their  coxae,"  instead  of  "as  long 
as" — a  careless  error. 

Ranatra  annuli'pes  Stal. 

Sfal,  Of.  Vet.  Akad.  Forh.,  XI,  p.  241,  1854. 

Original  description: 

R.  anmdipea :  Flavotestacea;  hemelytris  fuscescente  testaceis;  pedibus  obso- 
lete fuscoannulatis;  spiraculis  nigrofuscis.    Long.  30,  lat.  3%  millim. — Brasilia. 

In  1861,  under  the  title  "Genera  Nepidarum  synoptice  desposita," 
in  his  "Nova  methodus  familias  quasdam  Hemipterorum  dispon- 
endi,"  "Ofversigt  af  Kongl.  Vetenshaps  akademiens  forhandlingar, 
Arg.  18,  1861,  No.  4,  he  gives  a  more  satisfactory  description: 

Ranatra  annulipes.    Stal.  Nepidse,  1861,  p.  9. 

R.  annulipes.  Pallide  testaceo-grisea,  pedibus  obsolete  fusco  variis;  abdo- 
mine  dorso  sanguineo-fusco,  lateribus  griseo-flavescentibus,  stigmatibus  nigris, 


446  The  University  Science  Bulletin. 

dorso  pellucentibus ;  fronte  convexiuscula ;  alis  levissime  infuscatis;  prosterno 
bisulcato;  femoribus  anticis  pone  medium  intus  unidentatis,  extus  ibidem 
at  prope  apicem  inermibus,  intermediis  posticis  subbrevioribus,  his  ad  medium 
segmenti  ultimi  abdominis  porrigendis;  aidothecae  appendicibus  corpori  sequi- 
longis.    Long.  30  millim.    Brasilea  (Mus.  Hohn.). 

Ranaira  annulipes  Stal.  Of.  Vet.  Ak.  Forh.,  1854,  p.  241-1.  Prsecedentibus 
duabus  affinis  et  cum  iisdem  divisionem  forma  metasterni  distinctissimam, 
Americam  habitantem  formans.  Secundum  formam  metasterni  in  divisiones 
quattuor  distinctas,  determinationem  specierum  facilitantes  dividi  potest 
Ranatra  genus." 

Doctor  Montandon  (Bui.  Soc.  Sci.  Bucharest,  XVIII,  1910) ,  who 
has  examined  Guerin's  type  of  R.  fabricii  (1857)  pronounces  it  to 
be  the  same  as  R.  annulipes  Stal,  1854.  Sagra's  Historia  fisica,  po- 
litica  y  natural  de  la  Isla  de  Cuba,  volume  VII  (1857)  is  not  avail- 
able to  many  of  our  workers;  therefore,  Guerin-Meneville's  descrip- 
tion of  R.  fabricii  follows: 

Ranatra  fusca  tubo  respiratorio  corpora  paulo  longiore ;  pedibus  anteriori- 
bus  nigro-fuscis ;  corpoie  infra  obscure  feiTUgineo.    Larg.  35,  Anch.  3  millim. 

Notes.  This  species  is  distinguished  from  all  our  species  by  hav- 
ing the  connexivum  of  the  genital  segment  enlarged  and  extended 
ventrally  at  the  caudal  end  embracing  the  distal  portion  of  the  geni- 
tal operculum  (see  fig.  11,  pi.  XLV).  I  have  never  seen  specimens 
of  this  species  from  the  United  States  and  Prof.  H.  G.  Barber  in- 
forms me  that  his  record  from  Texas  was  on  a  mistaken  identifica- 
tion. Mr.  W.  L.  McAfee  says  the  species  appears  to  be  common 
in  Cuba. 

Ranatra  fusca  Palisot  Beauvois. 

Palisot  de  Beauvois,  A.  M.  F.  J.,  Ins.  Rec.  Afr.  Am.,  p.  235;    1805. 
(=  R.  Americana  Montd.,   1910,  and  subsequent  writers.) 

Original  description: 

RanaUre  brune.  Brune-verdatre ;  soies  un  peu  plus  courtes  que  le  corps; 
ailes  brunes-roligeatres.    (Fig.  1.) 

Ranatra  fusca.  V«ridi-f usca ;  setis  corpore  brevioribus;  alis  fusco-rubellis. 
(Fig.  1.)     Etats-Unis  d'Amerique. 

Obs.  Cette  espe^p  differe  de  celle  d'Europe  par  sa  couleur  plus  pale,  par  les 
soies  qui  terminent  I'abdomen,  plus  courtes  que  le  corps,  par  les  ailes  et  le 
dessus  de  I'abdomen  d'un  brun  rougeJitre. 

Notes.  A  photographic  reproduction  of  Palisot's  figure  (natural 
size)  is  given  on  plate  XLIX,  figure  1.  On  a  previous  page  I  have 
stated  the  evidence  to  show  that  Palisot's  illustration  is  natural  size. 
Size  alone  shows  that  Palisot's  species  is  not  the  same  as  R.  kir- 
kaldyi  Bueno.  It  also  eliminated  R.  nigra  H.  S.  {=  R.  protensa 
Montd.).    The  short  legs  and  stout  front  femora  eliminate  R.  fusca 


Hungerford:    Nepid.k  in  America.  447 

Bueno.  Even  after  making  due  allowances  for  "artist's  license," 
it  is  not  conceivable  tliat  the  careful  artist  illustrating  Palisot's  spe- 
cies could  have  been  looking  upon  R.  fusca  Bueno.  The  eyes  of  R. 
fiisca  Bueno  are  strikingly  large,  the  anterior  femora  very  slender, 
and  the  middle  and  hind  tibia?  much  larger  in  proportion  to  their 
femora  than  in  Ranatra  fusca  P.  B. 

Doctor  Montandon  (Bui.  Soc.  Sci.  Bucharest,  XIX,  1910),  who 
also  has  examined  Palisot's  figure,  says  in  effect  regarding  it:  ".  .  . 
When  one  considers  only  the  essential  details  for  the  characteristics 
of  a  form,  such  as  the  proportional  length  of  the  legs  and  append- 
ages, one  recognizes  without  difficulty  its  short  and  very  robust  an- 
terior femora,  its  very  little  developed  posterior  legs,  and  its  ap- 
pendages shorter  than  the  abdomen     .     .     ." 

He  was  led  into  error  by  assuming  the  figure  to  be  a  great  en- 
largement of  Bueno's  R.  kirkaldyi,  which  cannot  be  true  according 
to  the  facts  elsewhere  presented  in  this  paper.  Doctor  Montandon 
says  that  his  R.  americana  is  readily  distinguished  by  the  shape  of 
the  prothorax,  and  indeed  he  is  correct.  The  posterior  enlarged  part 
is  plump  and  then  narrows  to  a  slender  neck,  then  widens  again  in 
front.  Any  student  with  much  experience  with  American  Ranatra 
will  recognize  this  characteristic  at  once  (see  Palisot's  figure  repro- 
duced on  plate  XLIX,  figure  1).  Compare  it  with  the  photographs 
of  our  other  species.  Palisot's  insect  was  broad  and  large;  our  only 
species  comparable  to  it  is  R.  americana,  which  is  as  large,  some- 
times a  trifle  larger.  The  anterior  portion  of  prothorax  is  about 
one  and  one-half  times  the  swollen  posterior  portion;  so  it  is  in  R. 
americana  Montd.  (in  R.  fusca  Bueno  [=  R.  buenoi,  sp.  new]  the 
anterior  part  is  about  twice  the  posterior  part).  The  legs  are  short; 
so  with  R.  americana  Montd.  The  large  ratio  of  length  of  tarsus  to 
the  tibia,  1  to  5+  (in  R.  fusca  Bueno,  1  to  6+),  shows  similarity; 
also  the  hind  tibia  is  a  little  longer  than  the  femur,  but  not  as  much 
as  in  R.  fusca  Bueno.  The  broad  anterior  femora  agree  with  R. 
americana,  but  the  apical  tooth  is  not  shown.  This,  however,  is  a 
character  which  all  students  of  this  group  realize  could  be  overlooked 
readily.  Sometimes  the  space  in  front  of  the  tooth  is  filled  with 
debris,  obscuring  the  tooth;  sometimes  the  tooth  itself  is  much  re- 
duced. Doctor  Montandon  has  a  specimen  from  Philadelphia  and 
another  from  Texas  which  lack  the  tooth  and  which  he  designates 
under  the  name  R.  americana  var.  edentula.  In  many  of  the  speci- 
mens the  tooth,  therefore,  is  not  marked  and  must  have  been  over- 
looked by  Palisot's  artist.    The  caudal  filaments  are  comparatively 


448  The  University  Science  Bulletin. 

short  in  the  figure;  so  they  are  in  many  specimens  of  R.  americana. 
I  possess  examples  with  even  shorter  filaments.  On  the  whole,  I 
should  say  the  average  is  longer,  but  not  as  long  relatively  as  in  i?. 
buenoi,  sp.  new  (=  R.  fusca  Bueno),  or  in  R.  drakei,  sp.  new,  or  in 
R.  australis,  sp.  new.  It  is  most  robust  and,  on  the  whole,  our  larg- 
est species.  I  have  collected  it  in  New  York,  Minnesota  and  Kansas 
in  numbers,  and  have  seen  specimens  from  various  other  states. 

Ranatra  brevicollis  Montandon. 

Montandon,  A.  L.,  Bui.  Soc.  Sci.   Bucharest,  XVIII,  p.   184;    1910. 

Original  description: 

Ranatra  brcuicoliis  nov.  sp.  C'est  bien  a  regret  que  je  decris  cette  nouvelle 
forme  sur  un  exemplaire  malheureusement  unique,  assez  peu  dissemblable,  au 
premier  aspect  de  R.  quadridentata  Stal,  mais  ses  caracteres  specifiques  ne 
premettent  pas  de  la  confondre  avec  les  autres  especes  fusca  ou  quadridentata 
dont  elle  differe  par  ses  femures  anterieurs  tres  legerement  sinues  vers  leur 
extremite.  Elle  a  aussi  una  seule  Jent  devant  la  sinuosite  mediane  de  femur. 
Un  peu  plus  trapue,  de  fonne  moins  allongee  que  R.  jw<ca  Pall,  de  B.,  ce  qui 
pourrait  la  rapprocher  de  R.  quadridentata  Stal,  elle  se  separe  aussi  franche- 
ment  de  cette  derniere  par  la  forme  du  pronotum  beaucoup  plus  court.  En 
effet  I'insecte  a  34  mill,  de  longueur,  sur  lesquels  la  tete  et  le  pronotum  n'ont 
que  10  mill.  Les  appendices  de  22  mill,  sont  sensiblement  plus  courts  que 
I'abdomen.  Les  femurs  intermedif»res  et  posterieurs  courts,  replies  en  avant 
depassent  a  peine  la  tete. 

Le  pronotum  tres  robuste  presque  trois  fois  plus  court  que  Tabdomen,  assez 
forttment  dilate  en  a\-ant  et  tres  fortement  elargi  en  arriere,  ne  permet  pas  de 
la  confondre  avec  R.  kirkaldyi  T.  B.  II  est  en  outre  marque  de  deux  sillons 
longitudinaux  un  peu  obliques  sus  les  cotes,  derriere  la  dilatation  anterieure, 
n'atteignant  pas  en  avriere  les  sillons  transversaux  qui  limitent  en  avant  la 
partie  posterieure  dilatee  du  pronotum.  Cette  derniere  marquee  d'une  carene 
longitudinale  mediane  evanescente  en  arriere,  mieux  accentuee  en  avant  ou 
elle  traverse  les  sillons  transvei-so^ux  qui  limitent  la  partie  posterieure  dilatee. 

Les  pattes  pas  tres  greles,  un  peu  plus  courtes  proportionnellement  que 
celles  de  R.  quadridentata  Stal;  femurs  rougeatres,  marques  d"  anneaux  pales, 
larges,  peu  visibles. 

Metasternum  en  plaque,  termine  au  milieu  en  arriere  par  un  prolongement 
retreci  entre  les  hanches  posterieures,  paraissant  plus  releve  que  chez  R.  fusca 
P.  de  B.  et  R.  quadridentata  Stal,  mais  moins  cependant  que  chez  R.  fabricii 
Guer.  :=  annulipes  Stal. 

Cette  espece  se  distingue  encore  des  trois  autres  formes  connues  de  I'Amer- 
ique  du  Nord  par  I'opercule  genital  9  depassant  un  peu  sous  la  base  des  ap- 
pendices. Le  segment  ventral  qui  precede  I'opercule  genital  presque  droit  sur 
son  faite  longitudinal  tres  peu  convexe  avant  I'extremite. 

L'espace  interoculaire  convexe  entre  les  yeux,  mais  sans  trace  de  tubercule, 
a  peine  plus  large  qu'un  ceil.    Les  yeux  tres  legerement  transversaux. 

San  Diego,  Cal.    Coll.  Coquillett.    U.  S.  N.  M.,  Washington. 


Hungp:hfori):    Nepid.e  in  America.  449 

Notes.  I  have  examined  the  type  at  the  United  States  National 
Museum,  the  single  example  from  which  Doctor  Montandon  drew 
his  description.  It  is  a  female.  There  is  also  a  male  with  same  data, 
that  probably  was  taken  with  the  type,  and  in  addition  there  are 
two  females  from  Eldorado  county,  California.  One  of  them  bears 
this  note:  "Fife  of  this  insects  were  seen  feeding  on  one  grass- 
hopper which  got  into  the  water."  Another  specimen,  a  female, 
from  Lindscy,  Cal.,  taken  by  C.  Pemberton,  is  also  in  the  National 
Museum.  I  have  in  my  collection  some  specimens  from  Laguna 
Beach,  Cal.,  taken  by  C.  T.  Dodds.  To  me  the  most  striking  char- 
acter of  Ranatra  brevicollis  is  the  apparently  truncate  head;  the 
head  does  not  appear  broader  than  anterior  part  of  pronotum,  which 
is  thick  and  short.  The  anterior  lateral  prolongation  of  the  penulti- 
mate segment  of  the  antenna  is  short,  but  little  more  than  half  the 
ultimate. 

The  salient  characters,  as  assigned  to  this  species  by  Doctor  Mon- 
tandon, are:  Body  thick-set,  pronotum  short,  appendages  shorter 
than  abdomen,  the  legs  not  very  slender,  a  little  shorter  proportion- 
ally than  those  of  R.  quadridentata  Stal;  metasternal  plate  with  a 
middle  prolongation  between  the  posterior  coxae,  more  elevated  than 
in  R.  fusca  P.  B.  (=  R.  buenoi,  sp.  new),  and  R.  quadridentata  Stal, 
but  less,  however,  than  in  R.  fabricii  Guer.  =  annulipes  Stal.  He 
says  also  that  the  genital  opercule  extends  a  little  under  the  base  of 
the  appendages  and  the  ventral  segment  which  precedes  the  genital 
opercule  almost  straight  on  its  longitudinal  summit,  very  little  con- 
vex before  the  extremity.  I  find,  however,  that  the  male  operculum 
does  not  extend  under  the  base  of  the  appendages. 

All  of  the  insects  of  this  species  which  I  have  seen  came  from 
California.  (See  fig.  2,  pi.  XLVIII;'fig.  3,  pi.  XLV;  fig.  14,  pi. 
XLVH;fig.  3,pl.  L.I 

Ranatra  australis,  sp.  new. 

Size.  Smallest  specimen  in  our  series  measures  32  mm.  from  tip  of  beak 
to  tip  of  abdomen  with  a  respiratory  tube  27  mm.  long.  The  largest  specimen 
is  37  mm.  long  with  a  tube  30  mm.  long. 

Shape.  On  the  whole  a  slender  species  with  a  long  prothorax  and  long 
hind  femora. 

Structural  peculiarities.  Eyes  normal;  juga;  very  prominent,  more  ele- 
vated than  tylus,  a  characteri.stic  that  distinguishes  this  species;  antennae 
with  lateral  prolongation  of  penultimate  .segment  nearly  as  long  as  ultimate. 
Prothorax  slender,  sides  fairly  straight,  the  anterior  portion  measured  on  the 
median  dorsal  line  two  more  or  less  times  the  posterior  swollen  part.  Respira- 
tory filaments  quite  long,  a  little  less  than  length  of  the  insect.    The  dasper 


450  The  University  Science  Bulletin. 

of  the  male  genital  capsule  with  the  anteapical  prolongation  truncate  and 
short  and  well  separated  from  the  apical.  (See  fig.  3,  pi.  XLVI.)  Front 
femora  broad  without  apical  tooth.  Hind  femora  surpassing  the  middle  of  the 
penultimate  segment  of  the  body  and  the  hind  tarsus  reduced  to  one-sixth 
of  its  tibia.    (See  fig.  4,  pi.  XLVIII;  fig.  1,  pi.  L;  and  fig.  15,  pi.  XLVII.) 

Notes.    Described  from  the  following: 

Holotype:     Male,    Colorado    county,   Texas,    June    24,    1922;    Mrs.    Grace 

Wiley. 
Allotype:     Female,   Colorado   county,   Texas,   June  23,    1922;    Mrs.    Grace 

"  Wiley. 
Paratypes:    Five  males  and  four  females,  Colorado   county,  Texas,  June 
23  and  June  24,  1922;  Mrs.  Grace  Wiley. 
Nine  males  and  three  females,  Gainesville,  Fla.,  June,  1918;  C.  J.  Drake. 
One  female.  New  Orleans,  La.,  June,  1915. 
One  male,  McComb,  Miss.,  July  27,  1921. 
One  female,  Fayette,  Miss.,  July  23,  1921;  C.  J.  Drake. 
One  male.  Mound,  La.,;  J.  C.  Bradley  (teneral  specimen). 
One  male,  Calyell,  La.,  June  16,  1917;  H.  H.  Knight. 
One  male,  Kissimmee  Lake,  Florida;  A.  N.Resse. 

Total  of  twenty-nine  specimens  from  Texas,  Florida,  Mississippi 
and  Louisiana.  The  last  specimen,  belonging  to  the  U.  S.  National 
Museum,  was  labeled  R.  fusca  by  Doctor  Montandon  in  1909,  and 
bears  the  following  interesting  note:  "Alligator  flea,  water  dog,  said 
to  bite  or  sting  severely.  Swamp  east  of  Lake  Kissimmee,  Osceola 
county." 

The  holotype  and  allotype  and  two  paratypes  are  in  the  Uni- 
versity of  Kansas  collection;  paratypes  are  also  in  the  collection 
of  Dr.  C.  J.  Drake,  Ames,  Iowa;  the  United  States  National  Mu- 
seum, Washington,  D.  C;  J.  R.  de  la  Torre  Bueno,  and  the  collec- 
tions of  Mrs.  Grace  Wiley  and  of  the  author. 

This  species  is  smaller,  slenderer  and  longer  limbed  than  R.  fusca 
{=R.  americana  Montd.).  It  differs  also  in  the  following  particu- 
lars : 

1.  The  jugae  more  prominent  than  tylus — not  true  in  R.  fiisca  P.  B. 

2.  The  eyes  are  smaller  than  in  R.  fusca  P.  B. 

3.  The  pronotum  is  longer;  the  anterior  part  two  more  or  less,  times  the 
posterior  part,  whereas  in  R.  jusca  P.  B.  it  is  P/^  to  1%+. 

4.  The  sides  of  prothorax  more  nearly  parallel  and  posterior  swollen  part 
not  so  swollen. 

5.  The  hind  margin  of  pronotum  roundly  and  broadly  emarginate,  whereas 
in  R.  fusca  P.  B.  {=zR.  americana  Montd.)  the  emargination  is  deeper  and 
narrower. 

6.  The  two  depressions  on  the  scutellum  are  deep  and  pitlike,  while  in  R. 
fusca  P.  B.  they  are  shallow  and  broad. 


Huncjekfohd:    NEPin.ii  in  America.  451 

7.  The  hind  femora  longer,  surpassing  the  middle  of  the  penultimate  body 
segment,  often  almost  attaining  its  caudal  margin.  Femora  not  so  de- 
veloped in  R.  fusca. 

8.  Hind  tarus  of  R.  amtralis  one-sixth  or  less  of  the  tibia;  one-fifth  or  less 
in  R.  fusca  P.  R. 

9.  Metaxyphus  usually  longer. 

10.  The  female  operculum  angulate  on  its  ventral  line,  while  it  slopes  gradu- 
ally and  is  longer  in  R.  jusca  P.  B. 

11.  The  respiratory  filaments  are  relatively  longer  in  R.  austtralis  sp.  new,  than, 
in  R.  fusca  P.  B. 

12.  The  front  femora  lack  the  apical  tooth;  R.  fusca  P.  B.  has  one  more  or 
less  marked. 

Ranatra  drakei,  sp.  new. 

Size.  Length  from  tip  of  beak  to  tip  of  abdomen  35  mm.  to  46  mm.;  iu 
addition  to  this,  the  respiratory  filaments  are  from  28  mm.  to  44  mm.  long. 

Color.  All  the  specimens  in  the  series  studied  are  yellowish  brown  with  legs 
and  tegmena  overcast  with  an  orange  tinge. 

Shape.  A  long,  slender  species  with  prominent  eyes;  long,  slender  protho- 
rax;  hind  femora  surpassing  the  middle  of  the  last  abdominal  segment,  and  a 
ver>'  long  respiratory  tube. 

Structural  peculiarities.  The  eyes  very  prominent,  transverse  diameter 
greater  than  interocular  space;  jugse  prominent  and  divergent;  antennae 
with  lateral  prolongation  of  penultimate  segment  a  little  more  than  half  the 
length  of  the  ultimate  segment;  prothorax  slender,  the  anterior  portion 
measured  on  the  median  dorsal  line  2V^  times  length  of  the  posterior  swollen 
part.  The  length  of  abdomen  is  to  length  of  pronotum  as  2^/4  is  to  1;  the 
respiratory  filaments  long,  as  long  as  entire  body  in  many  of  the  specimens, 
greatly  surpassing  the  limbs.  The  claspers  of  the  male  are  shown  on  plate 
XLVI,  figure  11.  The  limbs  are  long  and  slender;  front  fenlora  slender,  median 
tooth  considerably  nearer  apex  than  base;  distal  tooth  well  marked  and 
located  at  some  distance  away  from  the  tibial  joint,  this  distance  being  about 
one-fourth  the  length  of  that  part  of  femur  lying  in  front  of  the  median 
tooth  (see  fig.  3,  pi.  XLYIII) ;  middle  and  hind  femora  long;  distal  end  of 
hind  femora  attaining,  or  nearly  attaining,  the  caudal  margin  of  the  penulti- 
mate abdominal  segment;  the  ratio  between  femora  and  tibia  not  quite  but 
nearly  as  great  as  in  R.  buenoi,  sp.  new.  See  plate  L,  photograph  4  of  para- 
type  specimen.) 

Notes.  Described  from  eleven  specimens,  seven  males  and  four 
females,  taken  at  Gainesville,  Fla.,  ten  of  them  by  Carl  Drake,  June, 
1918,  and  one  specimen  taken  March  18,  1915,  collector  unknown; 
holotype  in  collection  of  Carl  Drake,  allotype  in  University  of  Kan- 
sas collection,  paratypes  in  the  above  collections  and  in  that  of  the 
author. 

This  species  has  the  general  appearance  of  R.  buenoi,  due  to  the 
large  eyes  and  elongate,  slender  body.  It  differs  from  that  species, 
however,  in  the  front  femur  possessing  a  well-defined  apical  tooth; 


452  The  University  Science  Bulletin. 

in  more  prominent  jugse;  in  differently  formed  antennae;  in  its  longer 
thorax;  in  the  respiratory  filaments  greatly  surpassing  the  limbs 
when  extended  backwards,  and  in  the  differently  formed  claspers  of 
the  genital  capsule  of  the  male.  It  cannot  be  confused  with  any 
other  of  our  species. 

Ranatra  quadridentata  Stal. 

Stal,  Ofversigt  af  Kongl.  Vetenskaps  akademiens  forhandlingar,  Arg.  18,  18G1,  No.  4, 
p.  204. 

Original  description: 

Grisea,  pedibus  immaculatis;  abdominis  dorso  sanguineo;  oculis  modice 
prominiilis;  thorace  antice  leviter  ampliato,  subtus  bisulcato;  alis  levissime 
infuscateiA;  femoribus  anticis  subtus  pone  medium  et  prope  apicem,  licet  hie 
obsolete,  bidentatis,  intermediis  ]:)osticis  vi.x  aequilongis,  his  basim  segmenti 
penultimi  abdominis  vix  superantibus;  metasterno  ut  in  praecedente.  Long.  33 
to  36  milhm.    Mexico.    (Mus.  Hohn.)     Prsecedenti  affinis. 

The  "preceding"  is  R.  unidentata,  from  Rio  Janeiro,  and  concern- 
ing its  metasternum  he  says, 

"metasterno  retrorsum  fere  ad  apicem  coxarum  posticarum  producto,  seg- 
mentum  ventrale  primum  tegente,  uti  videtur  postice  trilobato,  lobis  continuis, 
elongatis,  medio  convexo,  lateraUbus  depressis,  subarcuatis." 

Notes.  Van  Duzee's  catalogue  completely  submerges  United 
States  records  of  this  species  under  R.  americana  Montd.*  This  is 
not  justified,  because  Doctor  Snow  in  his  list  (Trans.  Kans.  Acad. 
Sci.,  vol.  XX,  pt.  1,  p.  153,  1906)  was  not  writing  about  the  insect 
that  Doctor  Montandon  described  as  R.  americana.  I  have  before 
me  Doctor  Snow's  insect  from  San  Bernardino  Ranch.  Cochise 
county,  Arizona,  the  same  thing  from  Mexico,  and  a  series  from 
Texas.  These  insects  agree  splendidly  with  Doctor  Montandon's 
comparative  notes  on  Stal's  species,  cotypes  of  which  he  has  studied 
carefully. 

When  Doctor  Montandon  described  his  R.  americana  (Bui.  Soc. 
Sci.  Bucharest,  XIX,  p.  65,  1910)  he  gave  some  remarkably  clear 
notes  of  comparison  between  his  species  and  Stal's  R.  quadridentata. 
The  latter  is  not  so  robust,  the  eyes  smaller,  not  so  transverse,  the 
interocular  space  not  so  convex;  cheeks  not  so  elongate,  but  not 
applied  so  closely  against  the  tylus,  which  is  shorter,  making  the 
head  before  the  eyes  appear  shorter.  The  median  construction  of 
the  pronotum  is  not  nearly  so  marked  and  the  anterior  part  more 
cylindrical.  I  may  add  that  the  anterior  part  of  the  prothorax  is 
longer  proportionally,  the  legs  are  longer  and  metaxyphus  longer. 

*  Van  Duzee's  catalogue,  quadrinotato.  Doctor  F.  H.  Snow  published  a  list  of  the  in- 
sects taken  by  him  in  Arizona,  and  listed  R.  quadridentata  Stal  as  one  of  them. 


Hungerford:    Nepid.e  in  America.  453 

SUMMARY. 

The  Ncpidae  of  North  America  north  of  Mexico  now  include  a 
total  of  eleven  species  and  two  varieties,  distributed  as  follows: 
one  Nepa,  two  Curicta,  and  eight  Eanatra  species  with  two  varieties. 
This  count  omits  R.  annulipes  Stal,  which  probably  does  not  occur 
in  our  range.  The  naming  of  our  two  commonest  and  widespread 
species,  under  the  two  oldest  names  given,  is  as  conclusively  shown 
as  can.be  done  without  authentic  types.  We  have  decided  that  our 
large,  robust,  short-limbed  form,  with  broad  anterior  femora,  is  R. 
fusca  P.  B.,  and  that  our  slender-limbed  form,  with  uniformly  short 
respiratory  filaments,  is  R.  nigra  H.  S.  This,  most  unfortunately, 
reduces  to  synonymy  R.  americana  Montd.  and  R.  protensa  Mondt., 
names  by  which  these  species  have  been  well  known  to  us  for  the 
past  ten  years.  The  antennae  and  genital  claspers  of  the  males  have 
been  of  value — very  striking  and  satisfactory  characters  in  most  of 
the  species.  The  genital  capsules  themselves  are  of  little  value  sys- 
tematically in  most  of  the  Ranatra.  The  intromittent  organ  of  the 
male  has  a  more  or  less  definite  and  characteristic  shape  at  its  tip 
(compare  figures  6-1  and  8-1,  plate  XLVIj.  Care  has  been  taken 
to  examine  material  from  various  localities  in  order  to  fix  the  limits 
of  variation  of  these  characters. 


454  The  University  Science  Bulletin. 


PLATE  XLIV. 

■  Fig.  1.  Egg  of  Ranatra  jusca  P.  B.  (=i2.  americana  Montd.)  dissected 
from  a  water-soaked  dead  cat-tail  blade. 

Fig.  2.  Diagrammatic  drawing  of  Ranatra.  A,  apical  tooth  of  front  femur. 
M,  median  tooth  of  front  femur.  C,  front  coxa.  G,  prothoracic  transverse 
grooves  which  separate  the  "anterior  part  of  the  pronotum"  from  the  "swollen 
posterior  part."    S,  scutellum. 

Figs.  3  and  5.  Stridulating  device  of  Ranatra.  Figure  5  shows  ventral  view 
of  head  and  a  portion  of  prothorax.  The  rubbing  of  the  roughened  patch  {A) 
on  the  base  of  the  coxa  (C)  against  the  file  (B)  on  the  inside  edge  of  the  an- 
terior lateral  margin  of  the  prothorax  (P)  produces  the  chirping  or  squeaky 
noise.  D,  antennae.  Figure  3  gives  an  enlarged  lateral  view  of  base  of  coxa  and 
anterior  lateral  margin  of  prothorax  cleared  so  that  the  file  shows  through. 
Lettering  same  in  both  figures. 

Fig.  4.  Ranatra  eggs  in  situ  in  soft,  decaying  cat-tail  leaf,  a  portion  of 
which  has  been  removed  to  expose  the  eggs. 

Fig.  6.  Genus  Curicta.  Note  that  it  is  intermediate  in  shape  between 
Ranatra  and  JVepa.  Until  the  appearance  of  this  bulletin  only  a  single  speci- 
men was  recorded  from  the  United  States. 

Fig.  7.  Egg  of  Curicta  drakei,  sp.  new,  dissected  from  the  tissues  of  a  de- 
caying plant  stem,  where  only  the  crown  of  fifteen  filaments  was  exposed. 
Drawing  made  from  material  secured  by  Mrs.  Grace  Wiley  in  her  life  history 
notes  on  this  species. 

Fig.  8.  Egg  of  Nepa  apiculata  Uhl.  Note  the  eleven  filaments  which  remain 
exposed  above  the  surface  of  the  plant  in  which  the  eggs  are  inserted. 

Fig.  9.  Genus  Nepa.  Represented  in  the  United  States  by  a  single  species, 
Nepa  apiculata  Uhl.  This  species  varies  considerably  in  size,  but  no  constant 
structural  detail  has  been  found  to  indicate  that  the  variants  are  not  conspe- 
cific. 


Hun'gerford:    Nepid.e  ix  America. 


455 


PLATE  XLIV. 


456  The  University  Science  Bulletin. 


PLATE  XLV. 

Fig.  1.  Clasper  of  male  of  R.  elongata  Fab.,  determined  by  Doctor  Mon- 
tandon.    S,  subapical  tooth. 

Fig.  2.  Genital  capsule  of  male  of  R.  elongata  Fab.,  determined  by  Doctor 
Montandon.  Compare  figures  1  and  2  with  figures  3  and  4.  Note  the  differ- 
ences in  shape  of  claspers,  their  relative  lengths  as  shown  in  figures  2  and  4. 
The  intromittent  organ,  I,  is  also  often  characteristic  of  a  species,  but  not 
always,  and  subject  to  some  variation.  The  capsule  itself  in  Ranatra  is  of 
little  value. 

FiGS^  3  AND  4.  Clasper  and  capsule  of  male  R.  brevicollis  Montd.  from 
California. 

Fig.  5.  Inside  view  of  left  clasper  of  Curicta  drakei,  sp.  new.  Compare 
with  figure  8,  C.  howardii  Montd.  The  shape  of  the  clasper  near  the  base  is 
not  the  same  in  the  two  species. 

Fig.  6.    Male  genital  capsule  of  C.  drakei,  sp.  new,  viewed  from  right  side. 

Fig.  7.    Male  genital  capsule  of  C.  howardii  Montd. 

Fig.  8.    Inside  view  of  left  clasper  of  C.  hoxmrdii  Montd. 

Fig.  9.  Ventral  view  of  head  of  Ranatra  to  show  the  antennae,  which  lie 
hidden.  Sometimes  they  are  covered  with  debris,  which  should  be  scraped 
away.  It  is  often  wise  to  relax  the  specimen  and  draw  the  antenna  into  a 
more  exposed  position.  The  front  coxa  also  sometimes  obscin'es  the  view  and 
should  be  moved  while  specimen  is  relaxed. 

Fig.  10.  Lateral  view  of  caudal  end  of  abdomen  and  base  of  respiratory 
filaments  of  Ranatra  nigra  H.  S.  C,  connexivum.  R,  respiratorj'  filaments. 
0,  operculum  of  genital  segment,  considered  in  this  paper  as  last  abdominal 
segment.    P,  considered  in  this  paper  as  the  penultimate  abdominal  segment. 

Fig.  11.  Lateral  view  of  caudal  end  of  abdomen  and  base  of  resjiiratory 
filaments  of  Ranatra  jabricii  Guer.  from  Cuba.  Montandon  says  it  is  identical 
with  R.  annulipes  Stal  from  Brazil. 

Fig.  12.    Dorsal  view  of  head  of  Ranntra.    B,  beak.    ./.  jugum.     T.  tylus. 


Huxgekfukd:    Nepid.k  in  America. 


457 


PLATE  XLV. 


458  The  University  Science  Bulletin. 


PLATE  XLVI. 

Figs.  1  and  2.  Clasper  and  male  genital  capsule  of  Ranatra  buenol,  sp. 
new.  Note  the  reduced  subapical  tooth  of  the  clasper  and  the  spatulate  form 
of  the  tip. 

Figs.  3  and  4.  Ranatra  australis,  sp.  new.  Note  the  truncate  subapical 
tooth  of  the  clasper  and  its  distance  from  the  apical  one. 

Figs.  5  and  6.  Ranatiia  kirkaldyi  Bueno.  S,  subapical  tooth.  C,  clasper. 
I,  intromittent  organ.  This  species  is  clearly  quite  distinct  in  its  relationship 
from  the  others.    Note  the  shape  of  the  intromittent  organ  and  of  the  clasper. 

Figs.  7  and  8.  Ranatra  quadridentata  Stal.  I  am  not  able  to  find  any 
constant  differences  between  this  and  R.  jusca  P.  B.  {^R.  americana  Montd.) 
in  respect  to  the  male  genitalia.  The  male  of  the  latter  species  has,  on  the 
whole,  a  more  slender  clasper. 

Figs.  9  and  10.  Ranatra  brachyura  Horv.,  1879,  said  to  be  same  as  R. 
sordidula  Dohrn.,  1860.  From  Japan.  Confused  by  a  student  of  this  family 
with  R.  protensa  Montd.  and  so  labeled.    Compare  figures  9  and  13. 

Figs.  11  and  12.    Ranatra  drakei,  sp.  new. 

Figs.  13  and  14.  Ranatra  nigra  H.  S.  (^R.  protensa  Montd.).  The  pre- 
ceding species  and  this  one  have  claspers  which  are  broad  in  their  middle 
parts,  but  the  shape  at  the  tip  distinguishes  them. 

Figs.  15  and  16.  Ranatra  linearis  Fab.  From  Euroj^e.  Berlese.  in  his 
Gli  Insetti,  p.  323,  figures  this  species.  Note  that  the  difference  between  the 
clasper  of  this  species  and  that  of  our  American  species  is  more  apparent  than 
the  superficial  appearance  of  the  insects  when  side  by  side  in  the  cabinet. 


Hungerford:    Nepid^e  in  America. 


459 


PLATE  XLVI. 


460  The  University  Science  Bulletin. 


PLATE  XLVII. 

Fig.  1.  Ventral  view  of  male  genital  capsule  of  Nepa  apiculata  Uhler. 
J'rom  North  America. 

Fig.  2.  Lateral  view  of  clasper  of  above.  Compare  with  figure  4  of  N. 
cinerea  L.  of  Europe. 

Fig.  3.   Ventral  view  of  male  genital  capsule  of  Ncpa  cinerea  L. 

Fig.  4.    Clasper,  lateral  view  of  Nepa  cinerea  L. 

Fig.  5.    Antenna  of  Nepa  apiculata  Uhler.    From  New  York. 

Fig.  6.  Antenna  of  Nepa  cinerea  L.  From  Europe.  U,  ultimate  antennal 
segment.  P,  penultimate  antennal  segment.  L,  lateral  prolongation  of  penulti- 
mate segment.  The  splendid  antennal  and  genital  characters  separating  these 
two  species  of  Nepa  illustrate  the  taxonopic  value  of  these  hitherto  unused 
structures  in  the  Nepidse.  In  the  drawings  which  follow,  do  not  attach  sig- 
nificance to  the  basal  segment,  but  to  the  last  two  segments. 

Fig.  7.    Antenna  of  Curicta  drakei,  sp.  new. 

Fig.  8.    Antenna  of  Curicta  howardii  Montd. 

Figs.  9  .and  U.  Antennse  of  Rnnatra  kirkaldyi  Bueno.  Sometimes  the  ulti- 
mate segment  is  fused  with  the  preceding  segment  and  sometimes  entirely 
separate. 

Fig.  10.    Antenna  of  Ranatra  nigra  H.  S.  (^R.  protensa  Montd.). 

Fig.  12.    Antenna  of  Ranatra  buenoi,  sp.  new. 

Fig.  13.    Antenna  of  Ranatra  drakei,  sp.  new. 

Fig.  14.    Antenna  of  Ranatra  brevicollis  Montd. 

Fig.  15.    Antenna  of  Ranatra  australis,  sp.  new. 

Fig.  16.    Antenna  of  Ranatra  quadridentata  Stal. 

Fig.  17.    Antenna  of  Ranatra  fiisca  P.  B.  (=J?.  americana  Montd.). 


HuNGERFORn:    Nepid.k  in  America. 


461 


PLATE  XLVII. 


II 


462  The  University  Science  Bulletin. 


PLATE  XLVIII. 

Fig.  1.  Front  femur,  tibia  and  tarsus  of  Ranatra  buenai,  sp.  new.  Note  its 
very  slender  form  and  the  position  of  the  median  tooth. 

Fig.  2.  Ranatra  brcvicollis  Montd.  The  median  tooth  is  nearer  the  middle 
than  in  any  of  the  others.    The  tibia  is  relatively  longer. 

Fig.  3.  Ranatra  drakei,  sp.  new.  Note  the  slender  form  and  the  position  of 
the  aincal  tooth. 

Fig.  4.  Ranatra  australis,  sp.  new.  It  lacks  the  apical  tooth,  and  the  an- 
terior portion  of  the  femur  is  not  enlarged  toward  the  distal  end  as  in 
R.  jusca  P.  B. 

Fig.  5.    Ranatra  fiisca  P.  B.  (=  R.  ainericana  Montd.).   From  Ithaca,  N.  Y. 

Fig.  6.  Ventral  view  of  head  and  prothorax  of  R.  buenoi,  sp.  new.  To  show 
the  deep  longitudinal  trough.    Compare  with  figure  10,  which  lacks  it. 

Fig.  7.  Ranatra  kirkaldyi  Bueno.  Note  that  the  femur  is  not  constricted 
in  the  region  of  the  median  tooth.  Compare  with  R.  fusca  P.  B.,  figure  5. 
This  from  Kansas. 

Fig.  8.   Ranatra  nigra  H.  S.  {=:  R.  protcnsa  Montd.) 

Fig.  9.  Ranatra  kirkaldyi  Bueno  var.  hofjnianni  new.  Apical  tooth  or 
marked  sinuosity  present.    From  Minnesota. 


Hiingerford:    Nepidk  ix  America. 


463 


PLATE  XLVIIT. 


464  The .  University  Science  Bulletin. 


PLATE  XLIX. 

(All  photographs  natural  size.) 

Photo  1.  Photograph  of  Palisot's  figure  of  Ranatra  jusca,  reproduced  exact 
size. 

Photo  2.  Ranatra  kirkaldyi  Bueno.  Largest  specimen  in  a  long  series. 
A  female  from  Cherokee  county,  Kansas,  determined  by  Mr.  J.  R.  de  la 
Torre  Bueno.  This  is  what  Doctor  Montandon  has  mistaken  for  Ranatra  jusca 
P.  B.  Besides  size,  note  shape  of  front  femur.  Not  constricted  as  it  is  in 
figure  1. 

Photo  3.  Ranatra  jusca  P.B.  (=  R.  americana  Montd.)  Note  the  general 
resemblance  to  original  figure  by  Palisot  (photo  1  above),  the  shape  of  the 
thorax,  the  broad  anterior  femora,  the  size  of  the  insect,  the  reduced  hind 
femora,  etc.  For  other  figures  of  this  insect,  see  plate  LI,  photographs  3  and  7. 
Figure  7  is  the  only  one  to  show  the  apical  tooth  of  the  front  femur  plainly. 
The  writer  has  forms  with  the  respiratory  tube  of  various  lengths.  The  short- 
est perfectly  formed  tube  is  shown  on  plate  LI,  figure  3. 

Photo  4.    Ranatra  kirkaldye  Bueno  var.  hojjmanni,  var.  new. 

Photo  5.  Ranatra  bitenoi,  sp.  new  {=R.  jusca  Bueno).  Compare  with 
Palisot's  R.  jusca  (photo  1).  The  relative  proportions  of  the  front  femora,  the 
length  and  shape  of  thorax,  limbs  and  filaments.  The  long  tibise  and  reduced 
tarsi.    This  photograph  is  of  a  paratype. 


PT.ATI-:  XT.TX. 


\\ 


(465) 


PLATE  L. 

.         (All  photographs  natural  size.) 

Photo  1.  Ranatra  australis,  sp.  new.  Differs  from  R.  jusca  P.B.  by  shape 
of  jirothorax.  longer  respiratory  tube,  lack  of  apical  tooth  on  front  femora 
and  by  its  elevated  jugse,  as  well  as  marked  differences  in  the  genital  claspers 
of  the  male.     Southern  in  distribution.     Photograph   from   paratype. 

Photo  2.  Ranatra  nigra  H.  S.  (=  R.  protcnsa  Montd.)  This  insect  in  a  long 
series  fits  the  original  description  most  remarkably,  except  for  the  color,  which 
is  of  no  consequence  at  all;  "Size  2-2 Vs  inches  from  beak  to  tip  of  filaments. 
Filaments  %  inch  long;  limbs  long  and  slender."  Photo  5  is  another  speci- 
men— the  latter  from  Minnesota,  the  former  from  Kansas. 

Photo  3.  Ranatra  brevicollis  Montd.  From  California.  Note  the  short, 
broad  thorax  and  its  relation  to  the  size  of  the  head. 

Photo  4.  Ranatra  drakei,  sp.  new.  Slender  front  femora  with  apical  tooth, 
large  eyes,  long  thorax,  long  limbs,  and  Aery  long  respiratory  filaments. 
(Photograph  from  paratype.) 

Photo  5.    Ranatra  nigra  H.  S.  (^R.  proteiisa  Montd.) 

(466) 


PLATE  L. 


/ 


J 


/ 


(467) 


PLATE  LI. 

(All  photographs  natural  size.) 

Photo  1.  R.  quadridenlata  Stal.  Small  specimen  taken  by  Doctor  Snow- 
in  Arizona. 

Photo  2.  Curicta  howardii  Moutcl.  Photograph  from  male  belonging  to 
Professor  Barber.    Compared  with  type  at  Washington  by  Doctor  Drake. 

Photo  3.  Ranatra  fusca  P.B.  Specimen  from  Minnesota  with  very  short 
but  perfectly  formed  re.spiratory  filaments.    The  tijjs  are  normal. 

Photo  4.    Curicta  drakei,  sp.  new.     Male  holotype. 

Photo  5.  Nepa  cinerea  L.  Male  from  Europe.  Compare  with  A^cpa  api- 
culcUa  Uhl.  from  Minnesota,  photo  8.  then  note  structural  differences  figured 
on  plate  XLVII. 

Photo  6.    Curicta  drakci,  sp.  new.    Female  allotype. 

Photo  7.  Ranatra  fusca  P.B.  Shows  how  long  the  respiratory  filaments 
may  be  and  how  marked  the  apical  tooth  of  front  femur. 

Photo  8.    Nepa  apiculata  Uhl.    Female  from  Minnesota. 


(468) 


PLATE  LI. 


/ 


/ 


8 


(469) 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  19— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  19.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

A  Study  of  the  Relation  Between  Function  and  Growth 
IN  Body  Cells Montrose  T.  Burrows. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

0-4622 


THE  KANSAS  UNIVERSITY 

SCIENCE  BULLETIN 

Vol.  XIV.]  October,  1922.  [No.  19. 


A  Study  of  the  Relation  Between  Function  and  Growth 

in  Body  Cells. 

BY  MONTROSE  T.  BURROWS,  M.  D. 

The  Department  of  Surgery,  Washington  University  School  of  Medicine  and  the  Barnard  Free 

Skin  and  Cancer  Hospital,  St.  Louis,  Mo. 

IN  THE  BODY,  growth  is  a  definitely  regulated  act,  which  dur- 
ing development  takes  place  irregularly  in  one  part  and  then  in 
another.  At  maturity  it  ceases  in  most  tissues,  except  as  it  is  neces- 
sary to  replace  tissue  or  cells  lost  by  injury,  to  meet  the  demands 
of  an  increased  function  of  the  whole  or  a  part,  and  to  take  care  of 
the  general  wear  and  tear.  Function,  on  the  other  hand,  such  as 
heart-muscle  contraction,  is  something  that,  once  established,  goes 
on  continuously  throughout  the  life  of  the  individual. 

Growth  in  the  body  is,  therefore,  that  which  has  a  maturity  or 
has  limits,  while  function  (rhythmical  heart-muscle  contraction) 
has  none. 

These  two  processes  are  related  in  only  one  regard:  for  a  given 
amount  of  work  of  contraction  there  is  a  constant  size.  A  stimulus 
which  induces  an  increase  in  work  on  the  part  of  the  heart  induces 
also  an  increase  in  the  size  of  the  organ,  while  one  which  induces  a 
decrease  in  work  leads  to  atrophy.  What  is  true  for  the  heart  is  also 
true  for  muscles  in  general  and  for  the  kidneys,  liver  and  other  or- 
gans of  the  body.  Growth  of  the  muscle  fibers  and  cells  of  these  or- 
gans is  proportional,  therefore,  to  their  rate  of  activity  or  to  the  rate 
of  their  metabolism.  That  the  same  is  true  in  the  embryo,  even  be- 
fore differentiation  and  the  development  of  function,  has  been  clearly 
pointed  out  by  Child.*  ^  Child  again  points  out  that  with  each  de- 
crease in  growth  rate  there  is  a  decrease  in  the  rate  of  metabolism, 

*  Superior  figures  refer  to   numbered   paragraphs  of   the   bibliography   at   the   end   of   this 
article. 

(475) 


476  The  University  Science  Bulletin. 

and  that  this  metabolic  rate  increases  markedly  in  lower  forms  after 
starvation  or  when  they  are  regenerating  lost  parts. 

The  question  of  growth  regulation  may,  therefore,  be  temporarily 
set  aside  for  a  study  of  the  conditions  which  regulate  metabolism  in 
general  in  the  cell.  As  the  tables  for  growth  clearly  show,  the 
metabolic  rate  in  the  organism  is  never  constant  (Child  and  Thomp- 
son).^ In  early  embryonic  life  the  increment  is  high,  then  it  gradu- 
ally decreases,  with  irregularities,  up  to  the  death  of  the  individual. 
In  the  early  stages  of  development  the  blastomeres  are  simple,  ex- 
panding, growing  cells.  Later  they  differentiate  and  assume  a  func- 
tional state.  That  this  differentiation  and  the  development  of  func- 
tion is  brought  about  by  the  continuous  changing  environment  ef- 
fected by  the  growth  and  filling  up  of  cells  in  the  growing  individual 
has  been  clearly  indicated  by  many  already  well-known  studies  of 
experimental  embryology.^  What  we  have  not  known  are  (1)  the 
nature  of  those  conditions  which  bring  about  these  changes,  and  (2) 
the  importance  of  these  changes  for  the  life  of  the  organism  itself. 

In  later  life  we  have  learned  to  recognize  the  fact,  however,  that 
all  tissues  do  not  become  actively  functioning  cells.  A  part  of  the 
mesenchyme  changes  to  muscle  and  glands,  while  another  part  re- 
mains in  what  appears  to  be  a  more  primitive  state,  such  as  the  cells 
of  the  interstitial  tissue,  bone  and  cartilage.  The  same  is  true  for 
epithelium.  A  part  is  used  for  covering  and  protection  and  another 
part  becomes  functioning  glands.  For  the  sake  of  simplicity,  I  shall 
call  the  cells  w^iich  are  active  in  the  glands,  in  the  muscle,  etc.,  as 
functioning  cells,  in  contradistinction  to  those  of  the  connective  tis- 
sue, bone,  cartilage  and  the  epithelial  coverings,  the  nonfunctioning 
cells.  This  is  not  a  strict  use  of  the  terms,  but  the  latter  resemble 
more  closely  in  their  behavior  the  undifferentiated  cells  of  early  life. 

There  was  a  time  when  it  was  thought  that  all  life  processes  cen- 
tered within  the  cell.  At  the  present  time  there  is  some  question 
whether  this  is  true.  The  primary  changes  in  early  development  ap- 
pear to  be  a  rapid  swelling  of  the  mass,  a  rapid  inhibition  of  water 
and  oxygen,  a  rapid  elimination  of  CO2,  and  a  secondary  splitting 
of  the  whole  into  cells.  Cellular  growth,  division  and  differentiation 
are  not  primary  activities,  but  apparently  secondary  to  other  more 
formative  forces  or  stimuli.  Life  does  not  manifest  itself  the  same 
throughout  the  life  of  the  animal.  In  the  beginning  it  is  recognized 
chiefly  by  the  rapid  expansion  of  the  whole,  the  rapid  division  of 
cells  and  the  careful  building  of  its  various  parts.  This  period  of 
building  is  completed  in  man,  as  is  well  known,  within  ten  days  after 


Burrows:    Study  of  Body  Cells.  477 

birth.  Subsequent  to  this  time  no  new  oi-^an  or  parts  of  organs  are 
formed  except  the  hiying  down  of  the  nerve  sheaths.  All  subse- 
quent growth  is  merely  the  expansion  of  previously  formed  struc- 
tures. It  is  like  hypertropliy  and  hyperplasia,  as  they  result  from 
stimulation  of  the  adult  organs. 

With  the  completion  of  the  building  of  the  organs,  function  has 
made  its  appearance.  Growth  unlike  that  of  the  earlier  period  now 
runs  hand  in  hand  with  new  forms  of  work.  This  growth  continues 
to  maturity,  when  it  ceases,  except  as  it  is  to  take  care  of  the  wear 
and  tear,  so  to  speak,  and  to  play  a  part  in  certain  organs  and  tis- 
sues, such  as  the  bone  marrow,  the  sex  glands,  and  the  nail-  and 
hair-beds.  Atrophy  then  slowly  intervenes.  This  continues  to  the 
inevitable  death. 

The  general  nature  of  the  structure  and  the  metabolism  which 
leads  to  this  sequence  of  changes  in  the  organism  is  the  pertinent 
biological  question  to-day.  ChikU  in  1915  reviews  the  general  the- 
ories that  have  been  advanced  and  attempts  to  give  a  physicochemi- 
cal  explanation  of  the  process.  Child,  appreciating  the  relation 
between  the  slowing  of  growth  and  the  decrease  in  metabolism, 
makes  the  general  assumption,  applying  the  "law  of  mass  action," 
that  the  decrease  in  metabolism  may  be  explained  as  the  result  of 
the  gradual  accumulation  within  the  cell  of  an  insoluble  substratum. 
Since  differentiation  and  the  development  of  function  takes  place 
hand  in  hand  with  this  slowing  of  metabolism,  he  looks  upon  them 
also  in  the  same  light.  He  considers  the  changes  in  the  structure — 
differentiation — the  result  of  the  same  slow  deposit  of  the  same  sub- 
stratum. This  substratum  he  conceives  as  one  of  the  products  of 
the  metabolic  reaction.  The  reaction  then,  like  any  incomplete  re- 
versible physicochemical  reaction,  becomes  slowed  and  this  sub- 
stratum accumulates,  and  ceases  when  it  reaches  a  certain  con- 
centration.   Death  is  the  equilibrium  point  for  this  reaction. 

While  there  is  little  doubt  that  many  heterogeneous,  like  many 
homogeneous,  reacting  systems  reach  their  equilibrium  by  this  route, 
there  does  not  seem  to  be  sufficient  evidence  to  show  that  the  body  is 
exactly  of  this  kind.  Again,  all  such  systems  which  do  arrive  at 
such  an  equilibrium  are  first  put  together  by  some  external  force. 
In  nature  one  cannot  obtain  more  energy  from  any  system  than  what 
has  been  put  into  it.  This  is  the  law  of  the  conservation  of  energy. 
.Child  looks  upon  rejuvenation,  then,  as  the  result  of  the  removal  of 
this  substratum  and  of  dedifferentiation.  Differentiation  is  not,  there- 
fore, according  to  Child,  a  change  peculiar  to  a  definite  period,  but 


478  The  University  Science  Bulletin. 

rather  the  result  of  growth,  or  the  dynamic  state  of  the  cell.    Senes- 
cence is  an  inevitable  consequence  suffered  by  all  cells,  whether  they 
be  unicellular  organisms  or  cells  of  the  metazoa.    To  put  it  in  his 
own  words,  "Senescence  is  a  necessary  and  inevitable  feature  of 
growth  and  differentiation,  while  rejuvenescence  is  associated  with 
reduction."    Differentiation  he  defines  as  a  process  of  specialization. 
It  takes  place  in  the  cells  when  they  are  suffering  a  moderately  ac- 
tive metabolic  rate.     Decrease  in  this  rate  favors  differentiation. 
Increase  it  and  dedifferentiation  tends  to  occur.    Reduction  is  dif- 
ferent.   It  is  not  the  reverse  of  growth.    Growth  is  the  "accumula- 
tion of  certain  substances  formed  in  the  course  of  the  reaction  which 
are  physiologically  more  stable  than  other  substances  that  break 
down,  furnish  energy,  and  are  eliminated."    Reduction  takes  place 
when  the   breakdown   is   not  balanced   by   the   synthesis.     Using 
Child's  own  data,  I  see  no  reason  why  this  process  may  not  be 
formulated,  however,  in  a  much  more  definite  manner  and  in  a  man- 
ner more  consistent  with  not  only  other  natural  phenomena,  but  also 
with  the  picture  of  development  and  later  life  as  they  have  pre- 
sented themselves  through   morphological   studies.     In  the   early 
periods  of  development,  just  after  the  fertilization  of  the  egg,  the 
rate  of  metabolism  is  high.    The  work  performed  during  this  period 
is  the  building  of  a  heterogeneous  mass  out  of  a  previous  simple  egg 
cell.    This   process  of  building  is  completed  early.    The  metabolism 
at  the  time  of  completion  is  lower  than  at  previous  times,  but  it  is 
still  high.     Subsequent  to  this  time  this  metabolic  rate,  then,  de- 
clines progressively  to  death.    The  picture  at  the  beginning  is  there- 
fore entirely  different  from  the  later  one.    It  is  the  building  of  the 
machine,  which,  when  once  established,  slowly  runs  down.    The  pic- 
ture in  development  is  that  of  the  forcible  putting  together  of  parts, 
which  are  then  to  react  with  one  another  to  produce  work.    This  re- 
action continues  in  each  case  to  an  equilibrium  or  death,  like  any 
such  machine  built  by  man.    While  the  metabolic  rate  is  high  in  this 
machine  it  grows.    This  growth  ceases  (maturity)  when  this  meta- 
bolic rate  reaches  a  certain  minimum.    Then  atrophy  slowly  sets  in. 
Normal  death  is  not  the  accumulation,  but  the  gradual  using  up  of 
the  parts.     It  is  the  passing  of  heterogeneous  system  to  a  state  of 
equilibrium. 

Such  an  explanation  looks  upon  the  life  of  the  organism,  there- 
fore, as  the  result  of  some  unknown  force,  active  only  in  its  early 
period,  beginning  with  the  fertilization  and  ending  with  the  forma- 
tion of  the  last  organ  and  parts  of  organs.    This  force  disappears  at 


Burrows:   Study  of  Body  Cells.  479 

birth  or  thereabouts.  Subsequent  to  this  time  hfe  is  merely  an  ex- 
pression of  the  gra(hial  deterioration  of  the  previously  built  hetero- 
geneous system  or  the  interaction  of  its  various  parts,  which  follow 
tlie  law  for  the  disintegration  of  all  such  heterogeneous  systems. 
Life  of  the  animal  ends  with  the  establishment  of  an  equilibrium  be- 
tween the  parts  of  the  organization  which  produce  the  work  that 
really  constitutes  life.  The  energy  for  the  building  of  the  first 
period  is  derived,  therefore,  in  the  later  period. 

Systemic  or  general  death  is  nothing  more,  therefore,  than  the 
establishment  of  a  true  equilibrium,  or  a  breakdown  in  one  of  the 
essential  parts  of  the  body.  This  is  not  the  end,  however.  What 
we  have  been  talking  about  so  far  is  the  life  of  the  organism,  and 
not  that  of  the  cell.  The  end  of  the  individual  does  not  mean  the 
end  of  all  its  parts.  The  cells  are  not  dead  at  this  period.  They  are 
all  intact.  This  system  is  able  to  reproduce  itself.  The  normal 
death  of  the  whole  is  not  the  result  of  the  death  of  the  cells.  The 
cells  which  make  up  the  whole  are  destroyed,  rather,  by  the  process 
of  the  death  of  the  whole.  These  cells,  as  we  now  know,  go  on  for- 
ever under  the  proper  circumstances.  Out  of  one  of  them  in  the 
old,  the  egg  cell,  comes  a  new  individual.  Their  destruction  at  the 
death  of  the  individual  is  the  result  of  this  general  death.  Their 
destruction  is  the  result  alone  of  their  position.  Systemic  life  and 
systemic  death  are  something  different,  therefore,  from  elemental 
life  and  elemental  death.  The  first  has  limits;  the  other  may  have 
none.  The  question  is.  What  is  the  nature  of  this  system  which  can 
show  such  changes?  This  cannot  be  solved  by  a  study  of  the  animal 
as  a  whole.  It  cannot  be  solved  by  the  study  of  the  amoeba,  be- 
cause it  is  an  animal  itself.  It  must  be  solved  through  a  study  of 
those  cells  which  go  through  these  changes,  those  cells  which  find 
this  building  of  an  animal  and  the  ultimate  disintegration  of  this 
animal  their  normal  means  for  preserving  their  kind.  The  amoeba 
need  take  no  such  complicated  route  to  preserve  its  kind.  These 
cells  of  the  animal  must  take  this  course  in  order  that  they  survive 
and  that  the  proper  environment  be  prepared  for  their  reproduction. 
Child  has  ignored  this  fact  completely.  He  has  assumed  a  continu- 
ous dynamic  state  for  the  cell  without  any  proof  of  its  existence. 
He  has  again  assumed  that  the  life  of  the  whole  is  only  an  elabora- 
tion of  the  life  of  the  cell.  He  has  assumed  differentiation  to  be  a 
peculiarity  of  all  cellular  life.  He  has  assumed  the  protoplasm  of 
cell  to  have  structure  capable  of  performing  work.  The  best  bio- 
logical work  of  the  last  century  has  not  only  added  no  proof  for 


480  The  University  Science  Bulletin. 

these  general  assumptions  formulated  by  the  earlier  authors,  but 
quite  the  reverse,  it  has  spoken  against  them.  As  Bayliss  states, 
there  is  not  evidence  that  cells  are  necessarily  dynamic.  The  best 
morphological  studies  of  protoplasm  have  again  failed  to  show  struc- 
ture in  many  cells  other  than  nucleus  and  centrosome.  As  Wilson 
clearly  states,  cellular  growth,  division  and  differentiation  are  not 
primary  factors  in  development,  but  secondary  to  more  formative 
forces  or  stimuli.  In  the  simple  formulation  which  I  have  given  above 
it  is  possible  to  understand  how  the  organism  not  only  reproduces 
itself,  but  it  expresses  definite  need  for  the  differential  changes  as 
they  come  into  existence.  The  energy  for  the  primary  building  is 
acquired  from  the  old.  The  old  is  a  machine  not  different  in  prin- 
ciple from  other  machines  of  nature. 

As  I  shall  show  in  the  following  pages,  the  main  criticism  of 
Child's  theory  is  that  he  utilizes  theories  to  build  theories.  He  ac- 
cepts the  idea  of  the  cell  as  the  unit  of  manifested  life  of  the  organ- 
ism without  question.  Before  any  theory  of  life  and  death  is  justi- 
fiable of  acceptance  it  is  necessary  that  the  true  nature  of  the  struc- 
ture and  the  metabolism  of  the  cell  be  ascertained.  This  is  not  go- 
ing to  be  accomplished  by  morphological  and  chemical  methods 
alone,  nor  by  a  study  of  the  metabolism  of  the  whole,  but  by 
methods  which  allow  us  to  study  directly  each  of  the  fundamental 
manifestations  of  life,  such  as  growth,  division,  differentiation, 
migratory  movement,  etc.  The  tissue  culture  has  given  us  this 
opportunity.  In  support  of  the  above  contentions  it  is  of  interest  to 
report  here  some  general  analyses  so  far  carried  out  by  this  method. 

THE  CONNECTIVE-TISSUE  CELLS. 

As  is  well  known,  practically  all  previous  work  on  the  cell  in  re- 
gard to  the  nature  of  its  energy-producing  reactions  and  the  man- 
ner of  the  transformation  of  this  energy  has  been  based  upon  the 
idea  that  the  cell  is  a  highly  organized  body.  While  for  years  it  has 
been  assumed  that  amoeboid  movements  are  comparable  to  surface 
tension  changes  in  liquids,  all  theories  have  been  based  upon  the 
idea  that  these  amcsboid  movements  are  the  result  of  localized 
changes  in  surface  tension  resulting  from  some  unknown  organiza- 
tion residing  within  the  cell.  All  modern  physiochemical  methods 
have  failed,  however,  to  reveal  any  such  organization.  This  has  led 
one  and  then  another  to  assume  that  the  organization  is  either  the 
result  of  a  slow  diffusion  of  substance  in  the  colloids  of  the  cell 
(Wells) ^,  a  peculiarity  of  colloids  not  yet  organized,  or  to  invisil'le 
membranes  traversing  the  protoplasm. 


BiRRows:    Sttdy  of  Body  Cells.  481 

As  early  as  1913"  I  had  noted,  however,  that  the  movement  of  the 
cell  of  the  organism  is  not  one  which  is  governed  necessarily  by  such 
factors.^  These  cells  in  the  medium  of  the  cultures  move  always 
out  and  away  from  other  cells ;  and  in  further  studies  carried  on  over 
several  years,  I  have  continually  noted  that  the  large  number  of 
these  cells  show  no  change  in  contour  during  their  movement.  Their 
movements  are  not  amoeboid.  They  glide  along  like  bodies  carried 
by  some  external  force. 

The  picture  observed  in  these  cells  is  not  that  of  a  highly  organized 
body,  but  one  liberating  a  surface-tension-lowering  substance. 
Their  movements  are  like  those  of  cayenne-pepper  granules  dropped 
on  the  surface  of  water.  When  a  number  of  such  granules  are  scat- 
tered on  the  surface  of  the  water  they  shoot  apart.  This  moving 
apart  is  the  result  of  the  liberation  by  them  of  a  surface-tension- 
lowering  substance.  This  accumulates  in  greater  concentration  be- 
tween the  granules.  These  granules  are  pulled  apart,  therefore,  by 
the  greater  force  of  the  water  surface  without.'' 

While  the  connective-tissue  cells  take  the  same  course  outward, 
they  never  become  completely  dispersed.  These  cells  again  fail  to 
show  movement  on  the  surface  or  within  a  liquid  medium,  but  move 
only  and  show  evidence  of  metabolism  in  the  presence  of  the  fibrino- 
gen contained  in  the  blood  plasma  which  I  had  used  chiefly  as  a 
medium.  In  a  liquid  medium  these  cells  round  off  to  perfect  spheres. 
In  the  plasma  cultures  they  stick  tightly  to  the  fibrin  formed  in  the 
coagulation  of  the  plasma.  In  contact  with  these  fibrils  they  spread 
out  to  take  various  shapes.  These  shapes  are  always  peculiar  to  the 
surface  of  their  contacts.^" 

Not  only  the  character  of  the  movement  of  these  cells,  but  their 
general  effect  upon  the  clot,  further  indicates  that  they  liberate  such 
a  surface-tension-lowering  substance.  This  substance  differs  in  its 
physical  properties  from  that  liberated  by  the  granules  of  cayenne 
pepper,  however,  in  that  it  is  apparently  not  soluble  in  water,  but  it 
is  adsorbed  or  chemically  combined  with  the  fibrinogen.  The  cells 
not  only  stick  tightly  to  the  fibrin,  but  they  occasion  its  formation. 
When  a  fragment  of  connective  tissue  is  placed  in  a  drop  of  plasma 
it  occasions  first  a  gelation  of  the  whole  of  the  layer,  and  then  later  a 
true  coagulation.  This  gelation  commences  at  the  tissue  border  and 
spreads  rapidly,  to  invade  ciuickly  the  whole  of  a  large  area  of  the 
fluid  plasma.  After  a  considerable  latent  period,  the  coagulation,  the 
formation  of  fibrin  and  serum,  commences.  This  true  coagulation 
again  begins  at  the  tissue  border  and  spreads  slowly  outwards,  to 


482       '  The  University  Science  Bulletin. 

involve  after  many  days  a  small  area  about  the  fragment.^^  With 
this  second  coagulation  the  cells  appear.  They  are  in  close  contact 
with  the  fibrin  and  they  glide  out  just  behind  the  spreading  area  of 
change  in  the  jelly  like  clot.  This  movement  continues  only  so  long 
as  the  process  of  coagulation  proceeds.  The  cells  are  elongated 
spindles  closely  cemented  to  the  fibrin,  yet  capable  of  gliding  on  its 
surface.  With  the  completion  of  the  coagulation  their  movement 
ceases.  They  come  to  rest.  In  this  state  they  will  remain  ap- 
parently indefinitely  unless  fresh  fibrinogen  is  added.  When  this  is 
added  they  again  occasion  its  coagulation,  move  into  it,  and  again 
come  to  rest  when  the  coagulation  is  completed. 

The  whole  picture  of  activity  in  these  cells  is  that  which  can  be 
readily  interpreted  in  terms  of  the  liberation  by  them  of  some  sub- 
stance insoluble  in  body  fluids,  but  readily  adsorbed  or  chem- 
ically combined  with  fibrinogen.  The  combination  leads  to  the 
formation  of  fibrin.  The  movement  and  the  clinging  of  the  cell  to 
the  fibrin  indicates  further  that  this  substance  is  one  which  has 
strong  affinities  for  the  cell.  These  cells  do  not  crawl,  but  they 
glide,  and  are  held  firmly  to  the  fibrin.  Their  gliding  is  directly 
proportional  to  the  spread  of  the  coagulant.  Such  can  take  place 
only  in  presence  of  a  substance  which  is  strongly  attracted,  not  only 
by  the  fibrinogen,  but  also  by  the  cell.  Energy  production  in  these 
cells  is  centered,  therefore,  about  this  substance.  How  it  is  formed  in 
the  cells  becomes,  then,  a  problem  of  interest.  The  fact  that  the 
cells  come  to  what  appears  to  be  complete  rest  in  the  coagulated 
clot  indicates  that  it  is  the  product  of  the  one  reaction  taking  place 
within  them. 

For  several  years  it  has  been  evident  that  energy  production  in  the 
body  is  the  result  of  chemical  change.  It  is  further  recognized,  as 
Bayliss*^  clearly  points  out,  that  hydrogen  and  carbon  enter  chiefly 
into  this  reaction.  Nitrogen  in  the  body  is  used  largely  for  the  build- 
ing of  the  substratum,  proteins,  in  which  these  reactions  proceed. 
This  indicates  that  the  reactions  leading  to  protein  syntheses  are 
different  from  those  producing  energy. 

What  has  not  been  shown  is  the  nature  of  the  products  formed 
in  this  energy-producing  reaction.  It  has  been  thought  that  oxygen 
enters  directly  into  this  reaction,  and  at  one  time  it  was  thought 
that  the  products  formed  were  largely  carbon  dioxide  and  water. 
Recent  careful  experiments  have  indicated  that  this  is  not  true. 
Fletcher'-  has  shown  clearly  that  oxygen  does  not  enter  into  the 
contraction  phase  of  muscle,  but  is  adsorbed  largely  in  the  recovery 


BiRROAvs:    Study  of  Body  Cells.  483 

pcriocl.  Muscle  may  contract  for  a  time  without  oxygen.  J.  Loeb^^ 
has  found  that  oxygen  is  nceessary  to  preserve  «the  structure  of  the 
fertilized  egg.  These  experiments  of  J.  Loeb  also  indicate  that  the 
metabolic  reaction  may  proceed  without  oxygen.  That  the  same 
is  true  for  the  animal  colls  has  long  been  known  by  pathologists. 
Cut  off  the  blood  supply  from  any  part  and  it  undergoes  a  coagu- 
lative  necrosis.  These  observations  suggest  strongly,  therefore, 
that  oxygen  plays  a  necessary  but  probably  a  secondary  role  in  this 
reaction.  It  acts  to  remove  certain  products  of  the  primary  reaction 
rather  than  enter  into  it. 

All  chemical  reactions,  as  it  is  now  fully  appreciated,  are  governed 

not  only  by  the  concentration  of  reacting  substances,  but  also  by  the 

concentration  of  one  or  all  of  the  products  formed.    Equilibrium  for 

a  simple  reversible  reaction  is  expressed  in  the  following  formula: 

A  +  B  +  C  +  .     .     .  =  A'4-B'  +  C'  +  .     .     . 

In  the  animal  organism  growth  is  not  determined  by  food  any 
more  than  it  is  determined  by  oxygen,  but  by  other  unknown  factors. 
Bardeen  showed  that  planaria  regenerate  their  parts  when  starved  as 
when  they  are  fed.^"*  Morgan  found  that  salamanders  regenerate 
their  legs  as  rapidly  when  starved  as  when  fed.^^  The  difference  is 
tiiat  the  starved  animals  suffer  marked  emaciation  and  a  general 
atrophy  of  their  organs. 

The  above  observations  of  the  connective-tissue  cells  indicates 
clearly  the  existence  of  a  previously  unrecognized  product  of  cell 
metabolism  which  is  evidently  insoluble  in  liquid  medium,  but 
rapidly  adsorbed  and  chemically  combined  with  fibrinogen.  This 
substance,  which  I  shall  designate  as  "L,"  is  also  an  active  blood 
coagulant.  It  is  something,  therefore,  which  is  insoluble  in  water, 
but  capable  of  combining  with  fibrinogen  to  form  an  insoluble  com- 
bination, fibrin.  It  acts  in  every  regard,  therefore,  like  the  lipoid 
fraction  of  the  blood  coagulant  recognized  by  Woolbridge^^  and 
recently  carefully  studied  by  Mills. ^'^  This  fraction  is  a  phospholipin 
or  a  group  of  such  bodies. 

In  a  recent  series  of  experiments^  I  undertook,  therefore,  to 
ascertain  more  carefully  the  exact  relation  to  this  substance  of  the 
general  chemical  changes  of  these  cells.  These  cells  had  apparently 
become  inactive  after  they  had  come  to  rest  in  the  clotted  fibrin. 
They  had  laid  for  months  in  this  inactive  state  and  in  the  presence  of 
oxygen.  When  removed  to  fresh  medium  they  had  again  become 
actively  migrating  cells.  I  removed  the  oxygen  from  about  these 
cells  during  their  movement  and  also  after  they  had  come  to  rest. 


484  The  University  Science  Bulletin. 

While  they  are  migrating  and  the  coaguhition  is  taking  place  they 
disintegrate  in  the  absence  of  oxygen.  They  suffer  a  coagulative 
necrosis.  After  they  come  to  rest  they  suffer  no  change  in  the 
absence  of  gas.  In  the  same  way,  these  cells  suspended  in*  a  liquid 
are  unaffected  by  the  absence  of  oxygen.  In  a  liquid  they  do  not 
metabolize  any  more  than  in  the  presence  of  a  fully  formed  fibrin. 
Any  slight  stimulus  which  occasions  their  movement  occasions  also 
their  disintegration  in  an  atmosphere  of  nitrogen. 

There  seemed  little  doubt,  therefore,  that  in  the  identification  of 
this  "L"  substance  the  regulator  of  the  metabolism  of  these  cells 
had  been  found.  It  is  not  something  which  is  continuously  washed 
away  by  the  blood  stream,  but,  quite  to  the  contrary,  it  has  very 
specific  affinities,  -and  when  brought  into  contact  with  fibrinogen  it 
forms  an  insoluble  fibrin.  This  suggested  strongly,  therefore,  that 
continuous  activity  in  these  cells  must  depend  not  alone  on  food  and 
oxygen,  but  also  upon  very  special  conditions  for  removing  these 
primary  products  from  the  cell.^  The  only  plentiful  substance 
which  I  have  been  able  to  find  in  the  adult  body  capable  of  such 
removal  is  fibrinogen.  This  is  transferred  by  the  "L"  substance 
into  fibrin.  In  the  body  the  messenchyme  cells  of  older  embryos 
and  the  connective-tissue  cells  of  adults  lay  down  extracellular 
fibrils.  Hertzler^^  has  shown  very  definitely  that  fibrin  forms  the 
basic  proteins  of  these  extracellular  or  collagen  fibrils. ^^  These 
results  of  Hertzler  have  been  confirmed  by  Baitsell-^  and  myself.^^ 

These  observations  indicate  that  the  connective-tissue  cells  of  the 
body  are  not  in  any  sense  continuously  active.  Their  only  function 
is  the  production  of  extracellular  fibrils.  They  do  not  secrete  these 
fibers,  but  coagulate  certain  proteins  formed  elsewhere  in  the  organ- 
ism. The  increase  in  these  fibrils  is  alone  indicative  of  an  active 
metabolism  in  them.  The  dynamic  state  of  the  organism  is  in  no 
sense,  therefore,  an  elaboration  of  the  dynamic  state  of  these  cells. 
It  must  depend  on  other  conditions.  This  led  me  to  investigate  more 
carefully  the  process  of  rhythmical  contraction  as  it  is  seen  in  heart- 
muscle  cells. 

In  my  earlier  studies  of  the  tissue  culture  I  had  already  shown 
that  differentiation  in  heart  muscle  is  a  purely  reversible  phenome- 
non.^^ The  actively  contracting  heart-muscle  cells  are  derived  pri- 
marily from  the  undifferentiated  mesenchyme.  When  the  fragments 
of  this  tissue  are  brought  into  contact  with  the  plasmatic  medium  of 
the  culture  the  heart-muscle  cells  at  the  edge  of  the  fragment  which 
had  been  contracting  lose  this  property  at  once.    They  migrate  into 


Burrows:   Study  of  Body  Cells.  485 

the  medium  and  behave  exactly  hke  the  simple  undifferentiated 
mesenchyme  cells.  In  1912  I  ^^  showed,  however,  that  in  a  few 
cases  these  cells  which  migrate  into  the  medium  may  differentiate 
again  and  develop  rhythm.  When  this  takes  place  it  is  interesting 
to  note  that  the  cells  by  chance  alone  had  come  into  a  very  peculiar 
relation  with  the  medium,  and  again  it  is  interesting  to  note  that  this 
rhythm  never  develops  while  the  cells  are  migrating,  but  always 
after  such  migration  has  ceased  and  the  coagulation  process  is  com- 
plete. 

The  differentiation  of  the  heart-muscle  cells  in  the  outer  medium 
takes  place  very  infrequently.  In  the  large  majority  of  the  culture 
of  the  older  embryos  and  adults  the  clots  cling  tightly  to  the  frag- 
ment. The  cells  migrate  in  contact  with  the  surface  of  the  fibrin 
fibrils.  When  differentiation  takes  place  the  process  is  different. 
The  clot  loosens  and  contracts  away  in  mass.  If  the  ends  of  a  few 
cells  remain  attached  to  this  clot  they  may  become  stretched 
through  the  serum  cavity  between  the  surface  of  the  medium  and 
the  end  of  the  fibrin  fibrils,  or  between  the  fragments  and  the  end  of 
these  fibrils.  After  the  coagulation  is  complete,  these  cells,  and 
these  cells  alone,  develop  rhythm.  If  they  be  removed  from  those 
contacts  and  be  placed  in  the  outer  medium  in  contact  with  the 
surface  of  the  fibrin  they  stretch  out  again  and  behave  like  simple 
mesenchyme  or  connective  tissue  cells.^ 

By  these  observations  it  became  possible,  therefore,  to  clear  up 
the  difficulty  of  the  earlier  observations  of  the  connective-tissue 
cells.  Dynamic  states  in  the  organism,  such  as  the  heartbeat,  are 
not  a  property  of  the  cell,  but  that  of  a  peculiar  organization  of  the 
environment.  These  cells  may  produce  the  energy  for  the  work  of 
the  body,  but  there  is  no  evidence  that  the  transformation  of  this 
energy  into  work  is  the  product  of  a  cell  organization  in  the  case  of 
the  heartbeat  any  more  than  it  had  been  found  to  be  in  the  case  of 
cellular  migration. 

What  is  true,  for  the  connective-tissue  cells  I  find  also  to  be  true 
for  the  epithelial  cells  of  the  skin  and  many  of  the  glands.  The 
gland  cells  in  the  cultures  lose  the  form  peculiar  to  them  in  the  or- 
ganism. They  stretch  out,  like  the  skin  epithelium,  to  form  broad, 
thin  sheets  of  cells.  These  cells  cannot  metabolize  except  in  the 
presence  of  fibrinogen  or  a  similar  adsorbing  substance.  They  differ 
from  the  connective  tissue  in  that  they  later  destroy  this  substance 
through  certain  added  proteolytic  properties.  They  thus  depend 
wholly  on  the  fibrinogen  for  their  activity  in  the  culture,  but  they 
remain  together  and  form  no  extracellular  fibrils,  in  that  they  later 


486  The  University  Science  Bulletin. 

destroy  the  fibrin  which  is  formed.-^  In  Hquid  media  they  show  no 
evident  activity. 

While  these  experiments  are  interesting  for  the  understanding  of 
the  general  problems  of  the  genesis  of  connective  tissue,  their 
broader  significance  lies  in  the  new  view  they  present  of  life  in  the 
organism.  They  show  that  the  continuous  activity  consistent  with 
life  is  not  a  property  of  these  nonfunctioning  cells.  There  must  be 
long  periods  of  time  when  the  connective-tissue  cells  show  no  ac- 
tivity. Life  must  be,  therefore,  wholly  a  part  of  the  functioning 
systems,  the  glands,  the  muscles,  the  brains,  etc.  In  early  life,  how- 
ever, this  cannot  be  true.  The  whole  problem  of  life  in  the  early 
embryo  is  centered  about  an  excessive  metabolism  and  an  active 
proliferation  of  the  undifferentiated  cells  of  this  earlier  period. 

Many  years  ago  Hofmeister,  Sachs  and  De  Bary  had  already 
shown  that  cellular  proliferation  is  not  a  primary  factor  in  the 
growth  of  plants.  They  thought  that  the  mass  forms  in  growing 
plants  before  it  breaks  into  cells.  That  the  same  is  true  for  animals 
has  been  clearly  enunciated  by  Whitman,  Adam  Sedgwick,  E.  B. 
Wilson  and  others.  In  development,  cellular  growth,  division  and 
differentiation  are  not  primary  factors,  but  they  are  again  secondary 
to  other  more  formative  forces.  Driesch  looked  upon  this  force  as 
something  apart  from  nature  which  cannot  be  solved.  Others  have 
not  looked  at  it  in  this  sense.  They  have  considered  that  this  early 
development  is  the  result  of  certain  reactions  which  occasion  such 
a  primary  building.  Many  general  physiologists  have  attempted  to 
arrive  at  its  solution  through  the  study  of  colloidal  swelling.  It  is 
evident,  however,  that  such  study  cannot  attain  directly  to  this  goal. 
At  the  best  they  can  only  indicate  analogies.  It  occurred  to  me  to 
attempt  the  solution  by  another  method.  The  above  studies  were 
made  with  connective-tissue  cells  of  adults  and  the  mesenchyme 
cells  and  heart-muscle  cells  of  older  embryos.  No  careful  compari- 
son had  been  made  of  these  cells  with  those  of  the  younger  embryos. 
In  the  younger  embryos  it  is  well  known  that  the  mesenchyme  cells 
form  no  extracellular  fibrils,  but  grow  actively  to  form  a  cellular 
syncytium.  The  fibrils  appear  only  in  later  embryonic  and  adult 
life. 

THE  BEHAVIOR  OF  THE  CELLS  OF  THE  YOUNGER  EMBRYOS. 

As  I  have  cited  in  a  previous  article,  M.  R.  and  W.  H.  Lewis,  in 
1911,^^  noted  an  active  movement  and  growth  of  cells  in  liquid 
medium  such  as  simple  salt  solution.  This,  as  I  have  just  stated,  is 
not  true  of  the  adult  connective-tissue  cells.  Harrison^^  in  analyzing 


Burrows:    Study  of  Body  Cells.  487 

this  movement  noted  that  the  sells  never  moved  directly  outwards 
into  the  liquid  but  always  at  its  surfaces.  In  Harrison's  earlier 
studies  he  had  attempted  to  cultivate  fragments  of  the  neural  tube  of 
frog  embryos  in  hanging  drops  of  serum.  No  evident  activity  was 
observed.  Success  was  attained  only  when  these  fragments  were 
placed  in  lymph  which  clotted  about  them.  In  studying  the  move- 
ment of  the  cells  in  lymph  Harrison  then  noted  that  the  cells  moved 
always  in  contact  with  the  fibrin  fibrils,  and  in  his  later  analysis  of 
the  culture  of  the  Lewises  he  found  the  cells  migrating  always  in 
contact  with  the  surface  of  the  cover  glass  or  on  the  free  surface  of 
the  medium.  Harrison^^  termed  this  phenomenon,  as  L.  Loeb-'^  had 
done,  "stereotropism."  He  considered  these  cells  apparently  highly 
complex  systems  whose  mechanism  for  movement  is  regulated 
through  such  contacts.  Both  authors  thought  this  a  common  prop- 
erty of  all  cells. 

After  a  careful  study  of  the  movement  of  these  embryonic  cells  in 
liquid  medium.  I  noted,  however,  that  these  cultures  of  the  Lewises 
were  applicable  only  for  the  movement  and  growth  of  the  cells  of 
the  younger  embryos.  No  activity  is  seen  about  the  normal  frag- 
ments of  heart  muscle  or,  other  mesenchyme  cells  of  older  embryos 
at  the  surface  of  the  liquid  or  in  contact  with  any  solid.  As  I 
have  stated  above,  blood  plasma  or  fibrinogen  is  the  one  common 
substance  of  the  body  capable  of  stimulating  activity  in  them. 

From  these  facts  it  seems  evident,  therefore,  that  it  is  not  solids 
in  a  general  sense  that  are  necessary  for  the  movement  of  these 
cells,  but  specific  adsorbing  substances.  The  younger  embryonic 
cells  differ  from  the  older  ones  in  that  they  may  move  at  the  sur- 
face of  the  salt  solution  and  liquid  media  as  well  as  in  the  clot. 
The  older  cells  had  lost  this  property.  This  leads  me  then  to 
analyze  more  carefully  this  movement  at  the  surface  of  the  me- 
dium. I  measured  the  position  of  the  cells  moving  at  the  cover- 
glass  surface  with  the  micrometer  of  the  microscope.  I  found  that 
these  cells  were  not  in  contact  with  the  cover  glass,  as  Harrison, 
Lewis-s  and  L.  Loeb  had  thought,  but  that  they  lay  often  a  con- 
siderable and  measurable  distance  below  it.  Again,  previous  to  the 
movement  of  the  cells  on  the  surfaces  of  the  hanging  drop,  these 
surfaces  change.  They  became  covered  by  a  film  or  scum,  which 
made  them  appear  leathery.  The  cells  in  every  case  moved,  grew 
and  divided  in  this  film  of  material.  This  substance  is  not  the 
coagulating  substance  "L"  liberated  by  the  other  cells.  It  is  son  e- 
thing  new,  which  acts  evidently  in  place  of  and  antagonistically  to 


488  The  University  Science  Bulletin. 

the  fibrinogen.  This  substance  formed  early  in  the  life  of  the  cul- 
ture. It  spreads  rapidly  to  cover  the  whole  of  the  medium.  The 
cells  later  invade  it  as  they  invade  the  plasma.  It  differs  from  the 
fibrinogen  in  that  it  is  a  much  more  active  stimulant.  The  cells 
form  no  insoluble  compound  with  it.  They  grow  actively  for  a  time 
within  it  and  invariably  disintegrate.  In  the  plasma  cultures  of 
these  younger  cells,  a  part  of  the  cells  may  at  first  invade  the  clot. 
They  soon  leave  the  fibrinogen,  however,  for  the  surface  film  which 
has  stronger  affinities  for  them.  Single  cells  may  also  liberate  this 
substance. 

While  I  do  not  know  the  exact  nature  of  this  substance,  it  is 
interesting  to  note  here  that  this  substance  disappears  from  or  its 
action  is  inhibited  about  fragments  of  undifferentiated  mesenchyme 
and  the  heart-muscle  tissue  in  the  cultures  between  the  ninth  and 
fourteenth  day  of  the  incubator  life  of  the  chick  embryo.  This  is 
not  true,  however,  of  other  tissue.  I  have  identified  it  in  the  epider- 
mis and  the  liver  of  these  embryos  as  late  as  the  eighteenth  day. 
Felix  has  shown  the  new  tubules  developed  in  the  kidney  of  man  as 
late  as  ten  days  after  birth.  While  these  studies  are  yet  incom- 
plete, they  do  indicate  that  the  disappearance  or  the  inhibition  of 
the  action  of  this  substance  is  not  regular,  but  takes  place  at  irregu- 
lar times  in  different  tissue  of  these  higher  animals,  and  that  this 
disappearance  corresponds  to  the  forms  taken  by  the  developing 
embryo  and  adult. 

What  might  be  the  exact  chemistry  of  this  substance  remains  to 
be  seen.  It  seems  most  plausible,  however,  that  the  secrets  of  the 
building  of  the  body  will  be  found  in  the  physical  properties  of  it. 
These  physical  properties  are,  first,  a  strong  affinity  for  water, 
and,  second,  an  ability  to  stimulate  an  excessive  metabolism. 

From  the  above  observation  it  became  evident,  therefore,  that  the 
cells  of  the  body  are  not  in  a  continuous  state  of  activity.  The 
activity  of  early  life  is  dependent  completely  upon  the  synthesis  of 
a  substance  which  removes  the  "L"  substance  froiH  the  cells.  This 
substance  is  gradually  superseded  in  later  life  by  fibrinogen.  Fibrin- 
ogen combines  with  "L"  to  form  fibrin,  an  insoluble  compound.  In 
the  later  periods  life  centers  about  the  functioning  or  differentiated 
systems.  To  prove  this  fact  more  definitely  it  became  of  interest  to 
study  the  effect  of  a  flowing  stream  of  serum  upon  rhythmically  con- 
tracting cells  and  the  dedifferentiated  muscle  cells  and  the  cells  of 
connective  tissue. 


Burrows:    Study  of  Body  Cells.  489 

THE  CONTRACTING  HEART  MUSCLE  CELLS. 

The  isolated  contracting  heart-muscle  cells  which  occasionally 
develop  in  these  cultures  from  cells  migrating  from  the  fragments 
contract  with  a  rhythm  like  that  of  the  whole  hearts  or  fragments 
of  the  heart  transferred  to  the  cultures.  Since  they  occur  most 
infrequently,  contracting  fragments  have  been  used  chiefly  for  this 
study. 

The  rhythm  of  the  fragments  and  the  whole  hearts  of  younger 
embryos  may  never  cease  when  transferred  from  the  chick  embryo 
to  the  medium.  If  it  does  it  commences  at  once  again  as  soon  as 
the  temperature  for  it  is  restored.  During  the  first  few  minutes 
or  hours  this  rhythm  is  regular;  and  in  the  case  of  the  whole  hearts 
and  fragments  of  the  ventricle,  it  is  the  same  as  that  which  occurs 
normally  in  the  body.  After  a  short  time,  however,  it  becomes 
irregular.  There  are  periods  of  activity  followed  by  periods  of 
complete  rest.  These  periods  of  activity,  as  I  have  previously  de- 
scribed them,  are  ushered  in  by  rapid,  strong  contractions.  These 
contractions  gradually  decrease  in  amplitude  and  rate  to  the  period 
of  complete  rest.  After  a  short  rest  period  the  active  rhythm  again 
intervenes,  and  so  on;  their  irregular  rhythm  may  continue  for  as 
long  as  eight  or  nine  days  in  a  single  hanging  drop. 

This  slowing  of  the  rhythm  and  rest,  I  concluded,  was  due  to  the 
temporary  accumulation  of  waste  products,  the  temporary  lack  of 
nutrient  substances,  or  both.  After  these  waste  products  had  slowly 
diffused  away  and  nutrient  substances  had  moved  in,  the  heart 
became  active  again.  To  prove  this  I  placed  several  such  fragments 
in  a  specially  devised  culture  chamber.  This  chamber  was  arranged 
so  that  serum  could  be  made  to  flow  continuously  along  a  cotton 
wick,  the  fibers  of  which  transversed  the  layer  of  plasmatic  medium. 
By  this  means  it  was  possible  to  continuously  wash  the  medium 
about  the  fragment.  This  culture  has  been  called  the  ''wick  cul- 
ture."-^ In  such  cultures  the  rhythm  of  the  fragments  remains 
regular  during  the  time  the  serum  is  flowing,  and  this  regular  rhythm 
continues  often  for  many  days  or  until  the  protoplasm  of  the  cells 
is  otherwise  destroyed  by  infection,  etc.  A  careful  comparison  of 
the  irregularities  in  the  case  of  the  contracting  heart-muscle  cells 
with  those  of  the  migrating  ones  shows  interesting  differences.  In 
the  case  of  the  migrating  and  growing  cells  there  are  no  intermittent 
rest  periods.  Migration  commences  after  a  latent  period.  It  con- 
tinues actively  for  a  time;  then  gradually  ceases.  In  the  same  cul- 
ture there  is  no  second  recovery  period. 


490  The  University  Science  Bulletin. 

Again  it  is  interesting  to  note  that  the  changes  in  the  clot  are 
different  in  the  case  of  these  two  activities.  The  migrating  cells 
liberate  a  substance  which  occasions  a  coagulation  of  the  clot. 
About  the  contracting  fragments  from  which  no  cells  are  growing 
no  such  changes  are  seen  in  the  clot.  Contraction  in  the  isolated 
cells  commences  after  they  have  ceased  to  migrate  and  show  these 
changes. 

The  substance  or  substances  liberated  by  the  heart  which  pre- 
vents its  contraction  and  which  are  evidently  concerned  with  this 
act  of  contraction  are  soluble  substances.  They  can  be  washed 
away  with  serum.  The  evidence  gleaned  from  the  above  studies 
on  the  migrating  cells  indicates  that  the  substance  or  substances 
which  occasioned  their  migration  are  insoluble  in  the  medium.  They 
are  adsorbed  or  chemically  combined  with  the  fibrinogen  to  form 
fibrin.  The  substances  which  accumulated  to  stop  the  activity  of 
the  heart  is  not  the  "L"  substance  noted  above. 

For  the  testing  of  the  connective  tissue  and  undifferentiated  heart- 
muscle  cells  I  have  used  a  glass  culture  chamber  (plate  LII)  instead 
of  the  one  described  in  the  previous  paper.-''  Otherwise  the  technic 
was  the  same. 

During  the  course  of  the  study  of  the  contracting  heart-muscle 
■cells  in  1912  I  had  thought  that  the  migrating  cells  move  more  rapidly 
.against  the  stream  and  that  they  became  more  dispersed  in  these 
cultures  than  in  the  simple  hanging  drops  of  medium.  These  differ- 
-ences  were  observed,  however,  in  but  a  few  cultures.  Later  I  noticed 
that  the  thickness  of  the  layers  of  medium  also  affected  the  move- 
ment of  the  cells.  The  cells  migrating  from  1  mm.  thick  fragments, 
which  had  been  placed  near  the  edge  of  the  hanging  drop,  moved 
more  actively  in  the  thin  edge'^  than  in  the  thicker  parts  of  the  layer 
within.^  Regulating  these  conditions  in  the  "wick  cultures,"  I  found 
that  the  flowing  serum  in  no  way  affected  the  migrating  heart- 
muscle  cells,  nor  did  it  effect  in,  any  way  the  movement  of  the 
epithelial  and  connective-tissue  cells  in  general.  The  only  cells 
:affected  were  the  leucocytes  and  lymphocytes.  In  the  simple  hang- 
ing-drop cultures  it  takes  several  transplants  to  effect  a  complete  re- 
moval of  the  leucocytes  and  lymphocytes  from  fragments  of  bone 
marrow,  lymph  gland  and  spleen.  In  these  "wick  cultures"  I  found 
fragments  of  spleen  entirely  stripped  of  these  cells  after  seventy-two 
hours.  The  leucocytes  and  lymphocytes  had  accumulated  in  masses 
■£it  tangled  parts  of  the  wick. 

The  conditions  which  regulate  the  movement  of  these  latter  cells 


Burrows:    Study  of  Body  Cells.  491 

are  evidently  different  from  those  which  control  the  movements  of 
the  fixed  tissue  cells  of  the  body.  Migratory  movements  and  growth 
of  the  heart-muscle  cells  are  effected  by  a  substance  which  is  insol- 
uble in  circulating  body  fluids.  Growth  in  these  cells  is  therefore  a 
purely  physico-chemical  reaction  which  proceeds  to  a  condition  of 
static  equilibrium.  For  growth  to  take  place  it  is  necessary  that 
this  substance  or  these  substances  be  removed.  As  they  increase  all 
activity  ceases.  This  substance  or  substances  I  have  designated 
as  "L." 

DISCUSSION  AND  CONCLUSIONS. 

From  these  observations  there  seemed  little  doubt,  therefore,  that 
the  dynamic  state  of  the  organism  is  not  in  any  sense  an  indication 
of  a  similar  dynamic  state  in  the  cell.  It  is  a  product  of  the  organi- 
zation of  the  body.  In  early  life  the  dynamic  state  is  associated 
with  the  formation  of  a  substance  or  substances  which  combine  with 
or  otherwise  remove  the  "L"  substance  of  the  cell  and  occasion  an 
active  metabolism  within  them.  This  substance  has  strong  affinities 
for  water.  It  is  thus  directly  concerned  with  the  early  building  of 
the  organism.  It  stimulates  not  only  an  excessive  metabolism  in 
these  cells,  but  also  may  occasion  the  primary  swelling  of  the  mass. 
This  substance  disappears,  or  rather  it  ceases  to  be  recognizable,  in 
the  early  period  of  development.  Subsequent  to  this  period,  life  be- 
comes subservient  to  new  organizations.  The  new  organizations,  the 
general  nature  of  which  I  have  illustrated  in  the  study  of  the  rhyth- 
mical contractions  of  muscle  cells,  are  not  unique,  however,  for  the 
tissue.  They  are  peculiar  to  all  functioning  cells.  The  nerve  fibers 
are  stretched  between  the  brain  and  an  end  organ.  Adrian  has 
shown  that  the  "all  or  nothing"  law  holds  for  this  tissue  like  it  does 
for  the  heart.  The  same  is  true  for  the  glands.  The  secretory  cells 
are  cells  which  have  a  free  end  and  one  attached  to  a  basement 
membrane.  Stop  up  the  ducts  of  one  of  these  glands  and  these  cells 
undergo  atrophy.  Such  observations  are  wholly  in  line  with  the  gen- 
eral facts  which  are  Imown  concerning  development.  In  man  the 
kidney  may  form  tubules  up  to  the  tenth  day  after  birth.  Other 
organs  cease  their  progressive  growth  much  earlier.  No  careful 
studies  are  known  concerning  the  time  the  heart  cells  cease  to  divide. 
In  the  later  embryonic  period,  at  least,  all  growth  in  the  heart  is 
represented  by  an  increase  in  the  size  of  the  fibers  rather  than  an  in- 
crease in  the  cells.  In  the  kidneys  and  glands  it  manifests  itself  in 
a  dilation  and  increase  in  length  of  tubules  and  a  flattening  of  their 


492  The  University  Science  Bulletin. 

lining  cells.  In  the  wound  the  exudate  is  the  active  stimulus  for 
growth.  It  stimulates  metabolism  through  a  direct  affinity  of  the 
"L"  substance  for  the  fibrinogen.  "L"  combines  with  fibrinogen  to 
form  fibrin.  The  fibrin  then  becomes  the  extracellular  or  collagen 
fibrils.  The  cells  remain  active  and  move  towards  the  center  of  the 
exudate  until  the  reaction  is  satisfied.  The  end  of  activity  is  the 
completion  of  the  reaction.  It  is  the  scar  for  the  connective  tissue 
and  continuity,  and  the  limiting  membrane  for  the  epithelial  cells. 
Through  the  analysis  of  the  simple  act  of  changes  of  shape  and 
the  movement  of  the  connective-tissue  cells  it  has  been  possible  to 
show  that  these  acts  are  not  the  result  of  any  complex  cellular  or- 
ganization, but  a  simple  reaction  between  substances  in  the  environ- 
ment and  a  substance  of  the  cell.  The  movement  is  the  result  of  the 
cohesion  of  this  substance  for  the  cell  on  one  side  and  specific  sub- 
stances in  the  medium  on  the  other.  In  this  act  of  locomotion  the 
cell  supplies  the  energy;  the  mechanism  otherwise  resides  wholly  in 
the  environment.  Again,  in  other  studies  I  have  been  able  to  show 
that  the  syntheses  for  growth  are  not  a  part  of  these  energy-pro- 
ducing reactions,  but  they  are  separate  reactions.  The  connective- 
tissue  cells  in  the  adult  organism  are  widely  separated  in  a  mass  of 
fibrillar  substance.  These  cells  migrate  into  the  plasma,  but  they 
do  not  grow.  In  the  plasma  culture,  growth  is  peculiar  alone  to  the 
more  cellular  fragments  of  embryonic  tissue,  granulation  tissue  or 
sarcomata.  By, the  use  of  embryonic  extracts  it  is  possible  to  stim- 
ulate the  metabolism  of  these  connective-tissue  cells.  Under  these 
conditions  the  widely  separated  cells  of  the  adult  fragments  will 
grow  and  divide.  About  the  more  densely  cellular  fragments  such 
stimulation  is  not  necessary.  It  is  harmful.  It  leads  to  the  destruc- 
tion of  the  cell,  a  breaking  down  of  the  proteins  of  the  protoplasm. 
If  these  same  fragments  are  teased  apart  so  the  cells  become  more 
dispersed,  growth  ceases  again.  For  growth  to  take  place  it  is  neces- 
sary that  the  cells  be  either  crowded  or  excessively  stimulated.  The 
important  factor  for  this  reaction  is  evidently  the  concentration  of 
certain  products  of  their  metabolism.  The  concentrating  of  such 
products  may  be  induced  by  the  crowding  of  cells  or  increasing  the 
rate  of  their  production.  These  cells  liberate  not  only  the  "L"  sub- 
stance, but  also  CO2  and  H2O.  How  many  other  substances  are 
formed  when  the  reaction  is  proceeding  has  not  been  determined. 
Protein  synthesis  is  not  a  part  of  the  ordinary  metabolic  reaction  of 
the  cell.    It  is  secondary  reaction  depending  upon  and  obtaining  the 


Burrows:   Study  of  Body  Cells.  493 

energy  necessary  for  it  from  the  energy-producing  reaction  of  the 
cell.  The  extracellular  fibers  of  the  connective  tissue  are  not  secre- 
tions of  the  cells  any  more  than  bone  or  cartilage  are  of  this  origin.^ 
These  fibers  are  the  combination  of  proteins  formed  elsewhere  in  the 
body  and  the  ''L"  substance  of  these  cells. 

There  is  no  reason  to  believe,  therefore,  that  protein  synthesis  is 
a  part  of  the  metabolism  of  the  cell.  It  is  something  different.  It 
is  a  form  of  work  produced.  Upon  it  growth  depends.  Growth  has 
never  been  shown  to  be  of  a  simple  chemical  or  physical  nature. 
It  is  the  result  of  the  careful  utilization  of  energy.  It  takes  place 
against  the  forces  of  nature.  Protein  synthesis,  like  muscular  con- 
traction, is  only  a  form  of  work  peculiar  to  body  organizations  and 
not  cellular  organizations. 

In  1917,^*^  and  again  in  recent  experiments,^  I  have  shown  that 
the  development  of  ryhthmical  contraction  is  not  associated  with 
any  fundamental  change  in  the  cell.  This  is  a  property  peculiar  to 
any  of  the  mesenchyme  cells  of  early  embryonic  life.  It  occurs  in 
the  fragments  of  this  tissue  and  in  the  cells  which  migrate  from 
them.  Its  development  is  the  result  of  a  chance  relationship  of  the 
cells  to  medium.  This  relation  is  wholly  dependent  upon  the 
physical  peculiarities  of  the  coagulation  of  the  plasma  clot.  The 
isolated  cells  which  develop  rhythmical  contraction  are  those  cells 
which  become  stretched  through  a  serum  cavity  between  the  sur- 
face of  the  medium  or  a  cellular  tissue  fragment  and  the  end  of 
fully  formed  fibrin  fibrils.  No  contractions  develop  until  these 
fibrin  fibrils  are  fully  formed.  What  is  true  for  the  isolated  cell 
is  also  true  for  the  fragments.  Fragments  of  the  heart  of  young 
embryos  contract  at  once  when  removed  to  warm  medium  of 
the  culture.  Those  from  older  embryos  fail  to  show  this  change. 
Rhythm  develops  in  these  latter  fragments  only  after  the  border 
cells  have  moved  out  or  the  process  of  coagulation  is  completed. 
The  end  of  the  cell  in  contact  with  the  fibrin  is  in  metabolic  equi- 
librium. These  cells  imbedded  in  fully  formed  clot  cease  all  ac- 
tivity. The  cells  floating  in  serum  also  show  no  change.  The  only 
active  part  of  these  cells  differentiated  for  contraction  is  the  free 
end  in  contact  with  the  cellular  fragment  or  the  surface  of  the 
medium.  This  end  suffers  a  decrease  in  surface  tension.  Such  a 
decrease  in  surface  tension  is  associated  with  electrical  changes  at 
this  end  of  the  cell  and  a  stretching  of  the  cell.  If  such  changes 
continue,  one  of  three  conditions  must  result:  the  cell  will  be  torn 
loose  or  in  two,  or  there  will  be  an  explosive  breakdown  of  this 


494  The  University  Science  Bulletin. 

surface-tension-lowering  substance.  That  such  an  explosive  break- 
down is  peculiar  to  the  contraction  is  clearly  indicated  by  corre- 
lating these  observations  with  those  of  Fletcher.  Fletcher  has  found 
that  lactic  acid  is  liberated  during  the  contraction  phase  of  muscle. 
I  find  that  lactic  acid  increases  the  surface  tension  of  the  cells.  It 
causes  the  cell  to  contract.  This  lactic  acid  disappears  again  dur- 
ing the  intermittent  rest  period.  The  cell  again  returns  to  its  former 
state.  The  process  is  rhythmical.  The  process,  as  it  is  evident,  is 
somewhat  similar,  therefore,  to  the  phenomenon  described  by  Bredig. 
Bredig  showed  that  when  a  ten  per  cent  solution  of  H2O2  is  placed 
over  the  surface  of  pure  mercury,  a  film  of  mercury  peroxide  forms 
at  the  surface.  This  leads  to  electrical  changes  in  this  system. 
Under  appropriate  conditions  the  peroxidate  breaks  down  again 
to  mercury  and  oxygen.  Then  the  layer  of  peroxidate  reforms.  The 
process  is  repeatable  or  rhythmical.  Bredig  and  his  students  further 
find  that  many  conditions  which  alter  rhythmical  muscular  con- 
traction also  alters  the  activity  of  this  model.^^ 

Rhythmical  muscular  contraction  is  not,  therefore,  as  Bayliss  sug- 
gests, the  result  of  rhythmical  stimuli,  but  evidently  the  result  of 
an  explosive  breakdown  leading  to  alternate  changes,  not  only  in 
surface  tension,  but  in  the  electrical  conditions  at  opposite  ends  of 
the  cell.  Bernstein'^"  several  years  ago  had  given  definite  evidence 
to  show  that  the  energy  of  muscular  contraction  is  surface  energy. 
He  found  that  muscles  suffer  the  thermal  changes  peculiar  alone  to 
surface  energy. 

In  such  organization,  as  it  -is  well  recognized,  there  is  no  reason 
that  such  an  explosive  breakdown  should  always  occur.  Such  is 
possible  only  when  the  decrease  in  surface  tension  at  the  free  end 
or  the  electrical  changes  are  of  such  a  degree  to  allow  a  current  to 
pass  through  the  resistent  cell.  This  change  at  the  free  end  may 
cease  before  such  is  possible.  For  a  breakdown  to  take  place  other 
special  conditions  or  stimuli  must  be  present.  That  this  is  the  con- 
dition of  most  of  the  functioning  tissue  of  the  body  has  been  well 
proven  by  the  studies  of  tissue  autonomy.  In  man,  whether  any 
muscular  tissue,  other  than  the  nodes  of  the  auricle,  are  autonomous, 
like  the  cell  of  the  culture,  I  think  is  questionable.  At  the  same 
time  it  is  evident  that  any  functioning  tissue  may  develop  such  a 
condition.  Whether  this  is  what  has  happened  in  many  of  the 
nervous  affections  might  be  a  problem  worth  investigating.  It  is 
not  surprising,  however,  that  in  other  animals  the  autonomy  is 
centered  in  other  tissues.  Carlson^^  finds  that  the  rhythm  of 
the  heart  of  the  limulus  is  not  automatic,  but  centered  in  certain 


Burrows:    Study  of  Body  Cells.  495 

ganglion  cells.  In  men  I  think  it  is  very  questionable  whether  the 
nervous  system,  any  more  than  the  striated  muscle  or  gland  cells, 
respond  without  the  aid  of  external  conditions.  It  is  upon  this  fact 
that  coordination  depends. 

There  is  no  evidence  from  the  above  observations  that  the  cells 
themselves  undergo  any  absolute  changes  during  these  fundamental 
changes  in  the  body.  For  a  good  many  years  it  has  been  well 
known  that  bone  and  cartilage  are  products  of  a  given  organiza- 
tion of  the  part  and  not  a  product  of  the  cell.  It  has  been  shown, 
for  instance,  that  bone  will  develop  in  the  pelvis  of  the  kidney  if 
the  blood  vessels  to  that  organ  are  ligated.  Asami  and  W.  DocTi^^ 
repeated  these  experiments  in  the  laboratory  and  proved  completely 
the  existence  of  these  changes,  as  the  other  authors  had  shown.  In 
the  tissue  culture  the  cells  migrate  readily,  not  only  from  fragments 
of  interstitial  tissue,  tendons  and  fascia,  but  also  from  fragments 
of  bone  and  cartilage.  The  bone  cells  and  cartilage  cells  behave  in 
every  way  like  the  other  connective-tissue  cells.  The  bone  and 
cartilage  remains  behind  like  the  extracellular  fibrils.  In  the  same 
way  the  adult  muscle  fiber  will  not  react  in  the  plasma,  but  I  have 
seen  the  nucleus  and  sarcoplasm  migrate  out  and  leave  the  adult 
fiber  behind.  In  the  plasma  this  mass  forms  perfect  connective- 
tissue  cells.  These  cells  react  also  in  every  way  like  the  ordinary 
connective-tissue  cells. 

Such  simple  fluid  systems  may  not  only  suffer  changes  in  shape 
and  differential  changes  of  tension  at  various  points  on  their  sur- 
face, but  thej^  may  also  suffer  additions  and  probably  subtractions 
from  themselves.  In  the  body  differentiation  is  not  only  the  result 
of  mechanical  changes  initiated  and  controlled  by  the  environment, 
but  it  may  be  also  chemical  in  nature.  As  the  above  studies  of  the 
skin  epithelium  show,  these  cells  do  not  suffer  fundamentally  from 
the  connective-tissue  cells.  For  metabolism  to  take  place  within 
them  it  is  necessary  that  the  ''L"  substance  be  removed  from  them. 
This  "L"  substance  is  not  different  in  these  cells  from  that  of  the 
connective-tissue  cells.  These  cells  differ  from  the  connective  tissue 
in  that  they  also  contain  a  proteolytic  ferment.  The  pancreas  cells 
again  differ  in  that  they  also  contain  a  fat-splitting  ferment.  They 
often  fail  to  migrate  into  the  clot.  They  lead  rather  to  the  rapid 
splitting  of  the  fat  of  the  plasma  to  fatty  acid  crystals.  This  pre- 
vents also  the  migration  of  the  connective-tissue  cells  from  these 
fragments  into  the  medium.  So  in  each  case  the  various  tissues 
differ  as  they  contain  their  own  peculiar  added  products.    The  fun- 


496  The  University  Science  Bulletin. 

damental  reaction  of  the  fixed  tissues  otherwise  remains  the  same. 
The  only  exception  to  this  rule  is  to  be  found  in  the  wandering  cells. 
These  cells  have  not  been  observed  to  grow  in  the  cultures.  They 
have  also  lost  their  ability  to  cause  a  true  coagulation  of  the  plasma. 
They  occasion  the  gelation  of  the  clot  and  can  move  only  in  contact 
with  this  jelly  like  mass,  but  they  cause  no  fibrin  formation.  They 
owe  their  spherical  shape  to  their  inability  to  form  true  surfaces  in 
the  medium.  They  move  by  a  mechanism  different  from  the  fixed 
tissue.  They  can  invade  these  fixed  tissues  in  the  presence,  at  least, 
of  an  exudate.  When  this  disappears  they  tend  to  move  back  into 
the  lymphatics  and  blood  capillaries.  They  do  not  repel  strongly 
the  moving  fixed  tissues  like  the  fixed  tissues  repel  each  other.  They 
have  lost  the  property  to  form  the  "L"  substances.  I  say  they  have 
lost  it  because  it  is  present  in  the  mother  cells  from  which  the  wan- 
dering cells  arise. 

To  what  extent  this  chemical  differentiation  is  reversible  like  that 
peculiar  to -the  mechanical  form  I  have  not  definitely  determined. 
There  is  evidence  to  show  that  the  mesenchyme  cells  may  arise  from 
epithelial  cells  even  in  late  embryonic  life,  but  for  the  most  part 
these  epithelial  cells  maintain  their  chemical  peculiarities  for  a 
long  time  in  cancerous  growths  and  in  the  cultures.  I  have  seen 
heart-muscle  cells  assume  the  characteristics  of  large  mononuclear 
cells.  The  reverse  has  not,  however,  so  far  been  proven.  Again  1 
have  seen  liver  cells,  after  repeated  transplantation,  behave  in  part 
at  least  like  the  connective-tissue  cells.  There  is  no  reason  to  be- 
lieve, however,  that  such  chemical  dedifferentiation  may  not  occur 
and  maintain  in  the  proper  environment. 

The  cells  of  the  organism  are  not,  therefore,  highly  complex  sys- 
tems. They  are  not  equipped  to  lead  an  independent  existence. 
They  do  not  age.  They  have  no  organization  for  work.  They  pro- 
duce the  energy,  but  the  work  or  their  various  manifestations  of  life 
is  dependent  wholly  upon  external  conditions  about  them.  The 
only  form  of  work  depending  on  an  evident  internal  organization  is 
cell  division.  The  forces  active  in  the  process  center  about  the 
centrosomes.  There  is  no  evidence,  however,  that  activation  of  these 
centers,  or  even  their  formation,  is  controlled  from  within.  The 
centrosomes  develop  probably  alone  in  response  to  external  stimuli 
(see  Hertwig,  Meade,  Morgan,  J.  Loeb,  E.  B.  Wilson^  and  others). 

Energy  production  in  these  cells  is  again  wholly  dependent  upon 
organization,  or  the  presence  of  specific  substances  which  split  or 
otherwise  make  the  removal  of  the  "L"  substances  possible.    Their 


Burrows:    Study  of  Body  Cells.  497 

activity  in  early  life  is  dependent  upon  the  synthesis  of  a  substance 
which  combines  or  otherwise  removes  the  "L"  substance.  In  later 
life  it  is  dependent  upon  a  mechanical  differentiation.  This  form 
of  synthesis  in  later  embryonic  life  forms  a  substance  which  is 
fibrinogen  or  closely  akin  it.  This  forms  an  insoluble  compound 
with  "L"  rather  than  the  soaplike  substance  of  the  earlier  period. 
In  the  proper  environment  it  occasions,  through  its  coagulation,  the 
form  necessary  for  the  dynamic  state. 

By  these  observations  it  has  been  possible,  therefore,  for  the  first 
time  to  define  differentiation  in  other  than  morphological  terms, 
and  to  compare  function  with  growth.  The  growth  of  the  undiffer- 
entiated cells  of  early  life  is  the  result  of  a  special  synthesis.  The 
development  of  function  is  the  result  of  a  slightly  different  one. 
Differentiation  is  quite  different,  therefore,  from  what  it  has  been 
conceived  to  be. 

While  these  observations  reduce  growth  and  function  to  simple 
physicochemical  formulation,  they  give  no  hint  as  to  the  cause  for 
this  change  in  synthesis  from  early  to  late  life.  Any  substance 
which  removes  the  "L"  causes  an  immediate  loss  of  mechanical 
differentiation.  The  heart-muscle  cells  in  contact  with  the  fibrin 
become  simple  mesenchyme  cells.  Only  under  proper  mechanical 
conditions  can  they  redifferentiate.  What  is  true  for  the  heart  is 
true  for  the  glands.  In  the  cultures  the  gland  cells  stretch  to  form 
membrane  like  the  skin.    They  lose  the  form  necessary  for  function. 

These  facts,  again,  do  not  explain  chemical  differentiation.  In 
the  above  observation  I  have  also  not  discussed  all  the  deficiency  of 
these  cells.  Besides  the  lack  of  any  organization  for  work  and  any 
means  within  themselves  to  allow  the  energy-producing  reaction  to 
proceed  under  the  ordinary  conditions  in  nature,  they  are  also  bereft 
of  the  property  of  using  the  crude  material  of  nature  for  this  energy- 
producing  reaction.  While  energy  production  in  the  body  is  derived 
chiefly  from  H  and  C,  these  cells  cannot  use  the  sugar  carried  to 
them  by  the  blood  stream  without  intervention  of  substances  from 
the  pancreas.  The  organization  of  the  whole  or  certain  of  its  parts 
are  again  essential  for  another  of  their  important  needs. 

It  is  upon  this  last  deficiency  that  the  chemical  syntheses  peculiar 
to  differentiation  must  depend.  The  body  is  a  machine  operated  like 
any  other  machine  in  nature.  Its  metabolism  has  no  complexities, 
as  most  biologists  would  have  it.  It  is  a  machine  which  is  able  to 
produce  energy  and  transform  it  into  work.  It  has  been  possible  to 
locate  these  different  reactions.    The  energy  is  produced  in  the  cell; 


498  The  University  Science  Bulletin. 

its  transformation  is  under  the  control  of  the  environment.  Protein 
synthesis  is  in  no  way  to  be  confused  with  the  energy-producing  re- 
action. In  the  body,  anything  which  increases  metabolism  will  lead 
to  growth.  Increase  this  still  more  and  protein  destruction  results. 
This  accounts  for  the  appearance  of  split  products  of  protein  in 
muscle  which  has  suffered  excessive  stimulation  (see  Bayliss  for  lit- 
erature) and  the  destruction  of  the  tissue  by  strong  growth  stimuli 
such  as  X-ray,  radium,  coal  tar,  arsenic,  etc. 

Such  a  machine  can  build  itself  in  the  manner  that  it  does  because 
it  utilizes  the  products  of  its  energy-producing  reaction  for  its  build- 
ing. The  building  is  only  the  sequence  of  change  its  original  funda- 
mental structure  undergoes  to  produce  the  final  necessary  work — 
the  work  of  supplying  an  adequate  amount  of  fuel  for  a  certain  pe- 
riod of  life  and  supplying  the  egg  with  an  adequate  supply  of  this 
material  (yolk)  to  carry  it  through  the  early  period  of  the  develop- 
ment of  the  whole.  This  yolk  supply  decreases  progressively.  This 
decrease  leads  to  the  changing  character  of  the  syntheses. 

The  extracellular  deposits  which  I  have  described  above  as  im- 
portant for  the  organization  peculiar  to  the  dynamic  state  of  later 
life  are  evidently  the  substratum  recognized  by  Child.  They  are  the 
result  of  protein  syntheses  peculiar  to  the  organization  of  a  certain 
period  of  the  development  of  the  whole.  They  must  be  the  result 
not  only  of  mechanical  but  also  chemical  changes  in  the  environ- 
ment. The  picture  of  the  organization  as  it  is  seen  through  the  study 
of  the  cells  is  in  no  sense  the  picture  of  one  continuous  metabolic 
change.  It  is  the  primary  building  of  a  heterogenous  system  fol- 
lowed by  the  gradual  decline  of  this  system  to  a  state  of  static 
equilibrium.  Elemental  life  and  elemental  death  are  not  compar- 
able to  systemic  life  and  systemic  death.  The  body  is  the  necessary 
cycle  that  these  cells  may  preserve  their  kind.  For  the  active  growth 
of  the  early  period  of  the  development  the  cell  draws  upon  the  yolk 
or  the  mother  for  its  supply  of  those  substances  necessary  for  this 
growth.  The  disappearance  for  this  supply  of  material  is  the  ap- 
pearance of  the  second,  or  the  functioning,  period.  The  building  of 
this  period  is  completed  within  ten  days  after  birth  in  man,  except 
for  the  laying  down  of  the  nerve  sheath.  The  syntheses  peculiar  for 
this  building  are  the  result  of  the  disappearance  of  the  substances 
carried  in  the  yolk.  The  important  factors  for  differentiation  are 
not  to  be  found,  therefore,  in  the  primary  reactions  of  the  cells,  but 
in  the  deficiencies  of  these  cells.     The  important  deficiency  which 


Burrows:    Study  of  Body  Cells.  499 

has  to  do  with  differentiation  is  their  inability  to  utilize  the  crude 
food  materials  supplied  by  nature.  This  deduction  we  are  forced 
to  believe,  however,  not  only  from  the  above  observations,  but  from 
a  host  of  already  carefully  accumulated  facts.  If  the  first  two 
blastomeres  of  many  lower  forms  are  separated,  two  animals  de- 
velop. The  yolk  is  also  separated  by  this  process.  Each  animal  is 
but  one-half  the  size.  Gudernatsch  has  shown  that  tadpoles  fed 
upon  thyroid  differentiate  within  a  few  days,  while  thymus  feeding 
delays  this  process  at  least  for  a  very  long  time.  Very  large  tad- 
poles may  be  thus  developed.  The  ultimate  heterogenicity  thus  de- 
veloped is  essential  for  subsequent  life  and  for  reproduction.  The 
egg  cell  builds  a  new  system  like  the  old  (heredity),  in  that  it  has 
acquired  from  the  old  a  supply  of  those  substances  necessary  to 
carry  it  through  the  building  of  the  new  system,  which  is  again  ca- 
pable of  preparing  these  substances  from  the  crude  materials  avail- 
able in  nature  wdthout.  The  building  results  from  the  changing 
syntheses  which  result  from  the  gradual  decreasing  yolk  supply. 
The  form  of  the  building  is  dependent,  therefore,  primarily  on  the 
original  constitution  of  the  yolk.  This  is  again  dependent  abso- 
lutely on  the  nature  of  the  machine  which  produces  it.  Each  ani- 
mal must,  therefore,  in  each  case  reproduce  its  kind.  The  problem 
of  heredity  is  thus  reduced  to  pure  physics  and  chemistry. 

Death  in  such  a  system  may  result,  therefore,  from  the  destruc- 
tion of  essential  parts,  or  the  inevitable  equilibrium  of  those  forces 
which  maintain  the  heterogenicity.  This  does  not  mean  a  funda- 
mental change  in  the  cell.  The  cell  succumbs  as  the  result  of  this 
breakdown.  There  is  no  reason  why  any  of  these  cells  may  not 
grow  actively  again  if  the  organization  is  changed  about  it.  Differ- 
entiation is  both  chemical  and  mechanical  in  nature.  Chemical  dif- 
ferentiation does  not  effect  the  fundamental  energy-producing  reac- 
tion of  the  fixed  tissue  cells.  It  is  an  indication  of  the  chemical 
heterogenicity  of  the  mature  organism.  The  mechanical  differen- 
tiation essential  for  the  dynamic  state  is  reversible.  According  to 
the  above  formulation,  mechanical  differentiation  in  early  life  is  the 
result  of  the  removal  and  decrease  in  certain  materials  of  the  yolk. 
If  a  change  in  organization  suitable  for  active  growth  should  take 
place  in  the  organism,  no  such  differentiation  should  follow.  The 
cells  suffering  such  changes  must  continue  to  grow  as  long  as  the 
body  supplies  the  necessary  substances.  The  body  must  supply 
those  substances  necessai-y  for  active  growth  as  long  as  it  survives. 


500  The  University  Science  Bulletin. 

A  rapid  utilization  of  these  substances  must  lead  to  a  rapid  atrophy 
and  death,  according,  also,  to  the  formulation  given  above. 

Such  a  reorganization,  it  is  evident,  cannot  take  place  in  early  life 
except  under  the  influence  of  most  powerful  external  stimuli  or  de- 
velopmental defects.  It  becomes,  however,  more  possible  in  later 
life  when  the  normal  forces  which  maintain  the  normal  heterogenic- 
ity  are  waning.  Cancer  represents  such  a  form  of  active  growth. 
It  is  a  disease  peculiar  to  later  life.  It  occurs  earlier  in  connective- 
tissue  areas  than  in  epithelial  tissue.  The  connective-tissue  cells, 
according  to  the  above  observations,  lose  their  property  for  inde- 
pendent growth  earlier  than  the  epithelial  tissue. 

In  previous  publications  ^  I  have  noted  that  cancerous  tissue 
grows  like  that  of  younger  embryos  and  liberates  the  same  or  a 
similar  growth  stimulus.^^  In  the  body  cancer  may  result  from 
congenital  abnormalities,  such  as  pigmented  moles.  It  occurs  more 
frequently  in  certain  families  of  mice  (Maude  Slye).  It  follows  the 
continuous  application  of  many  growth  stimuli  or  substances  capa- 
ble of  effecting  such  a  reorganization.  In  man,  as  pathologists  agree, 
it  follows  most  frequently  upon  long-standing  chronic  inflamma- 
tion (Billroth).=^« 

As  has  long  been  fully  appreciated,  the  impediment  which  has 
stood  in  the  way  of  advance  in  cancer  has  been  the  inadequacy  of 
our  knowledge  of  the  cell  and  its  relation  to  the  whole.  It  has  been 
the  endeavor  of  the  cancer  laboratory  in  St.  Louis  to  attack  the 
problem  from  this  routg.  Cancer  is  not  a  parasitic  disease.  It  is  a 
disease  which  follows  after  long,  continuous  stimulation.  Before 
cancer  can  be  understood  it  is  necessary  that  irritability  and  stimu- 
lation be  reduced  to  simple  terms.  The  essential  conditions  for 
mechanical  dedifferentiation,  or  loss  of  function,  and  the  production 
of  an  active  growth  of  cells  like  that  of  cancer  is  the  presence  of 
substance  capable  of  removing  the  "L."  This  "L"  has  many  of  the 
properties  of  the  phospholipins  isolated  by  Mills.  These  phospholi- 
pins  are  soluble  in  many  lipoid  solvents,  and  especially  products  of 
coal  tar.  We  have  studied  the  action  of  coal  tar,  and  find  that  it  be- 
haves in  the  tissue  like  the  substance  liberated  by  the  cancerous 
tissue  and  the  tissue  of  young  embryo  which  is  able  to  combine  with 
or  otherwise  remove  the  "L"  from  the  cells.  Coal  tar  thus  attracts 
the  cells  to  it  and  effects  their  dedifferentiation.  At  first  it  occasions 
their  disintegration.  Later  it  becomes  a  less  active  solvent  and 
occasions  an  active  proliferation  of  these  cells.  It  is  capable  not 
only  of  producing  many  of  the  symptoms  of  cancer  by  itself,  but  of 


Burrows:    iStidv  of  Body  Cells.  601 

effecting  the  ivurganization  lu'cessary  tor  true  cancer  to  develop. 
Long  use  of  coal  tar  leads  to  tiie  de\'el()pment  of  cancer.^'^ 

What  I  have  hoped  to  present  in  this  paper  is  not  only  the  general 
picture  of  the  cell  the  tissue  culture  has  so  far  revealed,  but  methods 
by  which  the  further  details  of  these  general  problems  may  be  at- 
tacked. From  the  above  observations  it  is  evident  that  the  problems 
relative  to  life  are  not  to  be  solved  by  chemical  and  morphological 
methods,  but  by  the  application  of  mechanism.  Heredity,  as  we 
have  seen,  is  dependent  wholly  on  the  chemistry  of  the  system,  but 
the  importance  of  these  chemical  substances  will  not  be  understood 
until  the  mechanisms  peculiar  for  the  various  manifestations  of  life 
have  been  isolated,  their  parts  and  the  energy  relation  between  these 
parts  have  been  fully  determined.  Upon  this  latter  knowledge  rests 
also  the  ultimate  control  of  cancer.  Cancer,  as  it  has  been  seen,  does 
not  arise  from  embryonal  cells.  It  is  not  the  mere  displacement  of 
cell  (Ribbert) ,  but  a  specific  reorganization  of  parts.  Such  a  re- 
organization means  a  breaking  down  of  the  normal  relation  of  cells 
in  the  adult.  The  normal  heterogenicity  is  maintained  through  the 
fact  that  each  kind  of  cell  moves  through  the  liberation  of  the  same 
surface-tension-lowering  substance.  Each  tissue  repels  its  neighbor 
like  each  cell  repels  each  other  cell  of  the  same  or  of  another  kind. 
It  is  a  breaking  down  of  this  barrier  through  the  synthesis  of  a  new 
substance  which  conditions  new  surface  relations.  It  is  an  invasion 
of  connective  tissue  by  the  epithelium  (Thiersch  and  Remak) .  This 
synthesis  becomes  possible  through  the  specific  rearrangement  of 
cells.  Such  rearrangements  may  result  from  developmental  defects 
or  through  the  action  of  stimulating  substances.  In  the  body  not 
only  quality  but  quantit}^  is  an  absolute  factor.  Systemic  life  and 
elemental  life  are  not  comparable.  The  cells  do  not  age.  Differen- 
tiation is  not  an  age  phenomenon.  The  law  of  the  conservation  of 
energy  holds  for  the  body  like  it  holds  for  all  natural  phenomena. 
Advancement  in  biology  is  to  be  made  through  the  development  of 
methods  for  the  study  of  the  mechanics  of  these  systems.  Mor- 
phology and  chemistry  must  be  supplemented  by  these  methods.  All 
parts  of  these  machines  are  not  visible.  Rhythmical  heart-muscle 
contraction  is  not  a  specific  chemical  reaction.  It  is  peculiar  to  a 
large  number  of  animals.  These  animals  differ  chemically  from 
each  other.    A  watch  is  a  watch  whether  it  is  made  of  gold  or  silver. 

The  heart  contracts  as  the  result  of  a  special  arrangement  of  its 
parts  and  the  specific  dynamic  properties  of  these  parts,  and  not 
through  the  specific  chemical  constitution  of  the  particular  protein 
which  compose  its  parts. 


502  The  University  Science  Bulletin. 

BIBLIOGRAPHY. 

1.  Child,  Charles  Manning.  1915.  Senescence  and  Rejuvenescence.  The 
University  of  Chicago  Press,  Chicago. 

2.  Thompson,  D'Arcy  W.  1917.  Growth  and  Form.  University  Press,  Cam- 
bridge, London. 

3.  Wilson,  E.  B.  1906.  The  Cell  in  Development  and  Inheritance.  Second 
edition.    London  and  New  York. 

4  Huxley,  T.  H.  1863.  Review  of  the  Cell  Theoiy.  British  and  Foreign 
Med.  Cir.  Rev.,  vol.  XII. 

5.  Wells,  H.  Gideon.  1914.  Chemical  Pathology.  Second  addition.  Phila- 
delphia and  London. 

6.  Bayliss,  W.  M.    1915.    Principles  of  General  Physiology.    London. 

7.  Burrows,  M.  T.  1913.  The  Tissue  Culture  as  a  Physiological  Method. 
Trans,  of 'the  Congress  of  Am.  Phy.  and  Surg.,  vol.  IX,  pp.  77-90. 

8.  Burrows,  M.  T.  1913.  Tissue  Culture  in  vitro.  XVII  the  International 
Congress  of  Medicine,  General  Pathology  and  Pathological  Anatomy,  Lon- 
don, pp.  217-237. 

9.  Burrows,  M.  T.  Titles  Read  May,  1923,  to  Appear  in  Oct.,  1923,  Proc.  of 
the  Society  of  Exp.  Biology  and  Medicine,  vol.  XXI. 

10.  Burrows,  M.  T.  1911.  The  Growth  of  Tissue  of  the  Chick  Embryo  Out- 
side the  Animal  Body  with  Special  Reference  to  the  Nervous  System. 
Journal  of  Exp.  Zool.,  vol.  10,  pp.  63-83. 

11  Burrows  M.  T.  1916-17.  Some  Factors  Regulating  Growth.  Anat. 
Record,  vol.  11,  pp.  335-339. 

12.  Fletcher,  W.  M.  1902.  Tlie  Relation  of  Oxygen  to  the  Survival  Metab- 
olism of  Muscle.    Jour,  of  Physiol.,  vol.  28,  pp.  474-498. 

13.  LoEB,  Jaques.  1913.  Artificial  Parthenogenesis  and  Fertilization.  Chicago 
University  Press,  Chicago. 

14.  Morgan,  T.  H.    1901.    Regeneration.    New  York  and  London. 

15.  MoRG.\N,  T.  H.  1906.  The  Physiology  of  Regeneration.  Jour,  of  Exp. 
Zool.,  vol.  3,  pp.  457-500. 

16.  WooLBRiDGE,  T.  C.  1893.  On  the  Chemistry  of  the  Blood,  and  Other 
Scientific  Papers.  The  Crotian  Lecture  on  the  Coagulation  of  the  Blood. 
London. 

17  Mills,  C.  A.  1921.  Chemical  Nature  of  Tissue  Coagulins.  Jour,  of  Biol. 
Chem.,  vol.  XLVI,  pp.  135-165. 

18.  Hertzler,  a.  E.  1904.  Peritoneal  Adhesions,  Their  Cause  and  Prevention. 
Tr.  West.  Surg.  Assoc,  vol.  XIV,  p.  76;  Anat.  Rec.  1915,  vol.  IX,  p.  83. 

19.  Hertzler,  Arthur  E.    1919.    The  Peritoneum.    Vol.  1,  chap.  V.    St.  Louis. 
20    Baitsell  G.  a.    1916.    The  Origin  and  Structure  of  Fibrous  Tissue  Formed 

in  Wound  Healing.    Jour.  Exp.  Med.,  vol.  XXVIII,  pp.  739-756. 

21.  Burrows,  M.  T.  1915.  An  Attempted  Analysis  of  Growth.  Anat.  Record, 
vol.  9,  No.  11.  And,  The  Tissue  Culture  in  Cancer.  Proc.  Second  Pan- 
Amer.  Sci.  Congress,  Washington,  section  VIII,  part  2,  pp.  494-496. 

22.  Burrows,  M.  T.  1912.  Rythmische  Kontraktionen  der  isolierten  Herz- 
muskelzelle  ausserhalb  des  organismus  Mumchener  medizinischen.  Wo- 
chenschrift,  No.  27,  pp.  1-10;  and  Science,  N.  S.,  XXXVI,  pp.  90-92. 

23.  Burrows,  M.  T.  1913.  Wound  Heahng  in  vitro.  Proc.  of  the  N.  Y.  Path. 
Soc,  N.  S.,  vol.  XIII,  Nos.  5  and  6. 

24.  M.  R.  and  W.  H.  Lewis.  1911.  The  Cultivation  of  Tissues  from  Chick 
Embrvo  in  Solution  of  NaCl,  CaCl2,  KCl  and  NaHCOs-  Anat.  Rec,  vol. 
5,  pp.  277-294. 

25.  Harrison,  R.  G.  1911.  On  the  Stereotropism  of  Embryonic  Cells.  Science, 
N.  S.,  vol.  XXXIV,  pp.  279-281. 


Burrows:    Study  of  Body  Cells.  503 

26.  Harrison,  R.  G.  1914.  The  Reaction  of  Embryonic  Cells  of  Solid  Struc- 
tures.   Jour,  of  Exp.  Zool.,  vol.  17,  pp.  521-544. 

27.  LoEB,  Leo.  1922.  Agglutination  and  Tissue  Formation  Science  N.  S., 
vol.  LVI,  pp.  237-240. 

28.  Lewis.  W.  H.  1922.  Is  Mesenchyme  a  Syncytium?  Anat.  Record,  vol.  23, 
pp.  177-184. 

29.  Burrows,  M.  T.  1912.  A  Method  of  Furnishing  a  Continuous  Supply  of 
New  Medium  to  a  Tissue  Culture  in  vitro.  Anat.  Record,  vol.  6,  pp. 
141-144. 

30.  Burrows,  M.  T.  1917-'18.  A  Note  on  the  Mechanism  of  Heart-muscle 
Contraction.    Am.  Jour.  Physiology,  vol.  45,  pp.  556-557. 

31.  Bredig,  S.,  und  WEiNMA-iTiR,  J.  1903.  R,  periodische  Kontaktkatalyse. 
Zeitsch  fur  Phys.  Chem.,  B.  42,  S.  601-611.  And,  Bredig,  S.  1907.  Bio- 
Chem.    Zeitchr.,  B.  6,  S.  283. 

32.  Bernstein,  J.  1908.  Uber  die  Temperaturcoeffizienten  der  Muskelonergie. 
Pfluger's  Arch.,  B.  122,  S.  129-195. 

33.  Carlson,  A.  J.  1904.  The  Ner\^ous  Origin  of  the  Heartbeat  in  Limulus 
and  the  Nervous  Nature  of  Coordination  or  Conduction  in  the  Heart. 
Amer.  Jour.  Phys.,  vol.  12,  pp.  67-74. 

34.  AsAMi,  GoicHi,  .\ND  Dock,  William.  1920.  Experimental  Studies  on  Hy- 
perpastic  Bone  Formation.    Jour.  Exp.  Med.,  vol.  XXXII,  pp.  745-766. 

35.  Burrows,  M.  T.  1923.  The  Experimental  Production  of  Malignant  Ulcers 
in  the  Rat.    Mo.  State  Med.  Jour.,  vol.  XX,  pp.  145-147. 

36.  EwiNG,  J.    1919.    Neoplastic  Diseases.    Philadelphia  and  London. 

37.  L.  H.  Jorstad.  A  Study  of  the  Behavior  of  Coal  Tar  in  the  Tissues.  Proc. 
of  the  Soc.  Exp.  Biol,  and  Med.,  Oct.  1923. 


504 


The  University  Science  Bulletin. 


PLATE  LII. 


The  "wick  culture"  chamber. 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  20— October,  1922. 

(Whole  Series,  Vol.   XXIV,  No.   20.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

Notes  on  the  Biology  of  Curicta  (Heteroptera), 

Grace  Olive   Wiley. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE.  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second -clas.s  matter. 

9-4522 


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SCIENCE  BULLETIN 

Vol.  XIV.  I  October,  1922.  [No.  20. 


Some  Notes  on  the  Biology  of  Curicta*  from  Texas. 

BY  GRACE  OLIVE  WILEY. 

BEHAVIOR  OF  ADULTS  AND  NYMPHS  OF  CURICTA. 

I  COLLECTED  several  pairs  of  adult  Curicta  and  placed  them  all 
in  one  glass  to  take  home  alive.  When  I  reached  home  most  of 
them  were  mating.  These  were  removed  and  placed  in  separate 
glasses,  and  remained  paired  for  several  hours. 

In  mating,  the  male  takes  a  position  to  one  side  of  the  female,  and 
usually  to  the  right.  If  to  the  right,  he  hooks  his  left  anterior  tarsus 
over  her  head;  if  to  the  left,  the  right  fore  tarsus  is  used. 

Both  nymphs  and  adults  seem  fond  of  getting  out  of  the  water 
and  lying  close  to  the  ground,  where  they  are  hardly  discernible.  I 
have  found  adults  almost  a  foot  from  the  water's  edge,  in  tangled 
plant  roots  and  under  rotten  pieces  of  wood.  Search  for  eggs  laid 
in  nature  provided  fruitless.  One  pair  was  mating.  I  am  half  in- 
clined to  believe  the  eggs  are  laid  in  soft  mud.f 

The  nymphs  are  very  agreeable,  in  that  they  do  not  feed  upon 
others  of  their  kind,  even  when  hungry.  They  like  small  notonec- 
tids,  corixids,  small  carabids,  fresh-water  shrimp,  and  such.  They 
refused  small  minnows,  however.  It  is  not  uncommon  to  see  three 
feeding  quietly  on  one  shrimp,  or  two  feeding  on  one  small  beetle. 
They  are  very  fond  of  mosquito  larvae. 

EGGS. 

Size.  About  1.75  mm.  long;  width  a  trifle  more  than  .75  mm.;  diameter  of 
crown  a  little  less  than  %  mm.  Rosette  of  filaments  at  tip,  numbering  15; 
length  of  filaments  almost  1  mm. 

*  Curicta  drakei  Hungerford. 

t  Have  now  found  the  eggs  deposited  in  the  tissue  of  dead  plant  stems  with  only  the- 
crown  of  filaments  visible. 

(507)  ,    .•^"^  '■ 


508  The  University  Science  Bulletin. 

Shape.  Elongate  oval,  one  end  slightly  tapering  and  rounded,  the  other 
smaller,  sloping  somewhat  obliquely  and  bearing  a  crown  or  rosette  of  fifteen 
long  filaments. 

Color.  Creamy  white,  with  filaments  somewhat  yellowish,  center  of  crown 
darker. 

Seven  eggs  were  laid  by  one  female  twelve  days  after  mating. 
These  were  not  inserted  in  soft  wood  or  in  plant  tissues,  althougii 
both  were  available.  The  female  was  lying  close  to  the  wet  sand 
and  the  eggs  were  laid  on  their  sides  on  the  sand  without  any  regu- 
larity or  order.  The  next  day  there  were  three  more  eggs,  and  in 
three  more  days  I  found  six  eggs  hidden  among  the  roots  of  water 
plants  and  in  slimy  accumulations  in  crotch  of  a  dead  twig. 

DESCRIPTION  OF  FIRST-INSTAR  NYMPH. 

Form  elongate  oval;  very  much  like  adult,  only  broader  in  proportion  to 
length.  Head  large,  much  narrower  than  prothorax  and  as  wide  as  long,  ex- 
cluding the  rostrum.    Eyes  globular,  small. 

Anterior  femur  quite  robust  and  armed  with  a  single  median  tooth,  plainly 
\isible  and  closer  to  the  base  than  the  apex  of  the  femur. 

Anterior  coxae  about  half  the  length  of  their  femora,  and  very  robust.  In- 
termediate and  posterior  legs  short.    One-segmented  tarsi. 

Color  when  first  hatched  pale  straw  yellow,  becoming  darker  with  brownish- 
black  markings.  Number  of  days  of  first  instar,  twelve.  Entire  length  of  in- 
sect from  tip  of  beak  to  end  of  respiratory  tube,  5  mm.;  width  across  abdo- 
men at  widest  part,  1.50  mm.;  width  across  eyes,  .75  mm.;  width  of  shoulders 
at  base  of  head,  1.15  mm.;  length  of  prothorax  on  median  line,  1  mm.;  length 
of  respiratory  tube,  a  trifle  more  than  .50  mm.;  length  of  anterior  femora,  1.40 
mm.;  length  of  anterior  tibiae,  .70  mm.;  length  of  anterior  tarsi  about  .25  mm.; 
length  of  anterior  coxae,  .70  mm.;  length  of  intermediate  femora,  1.40  mm.; 
length  of  intermediate  tibiae,  1.00  mm.;  length  of  intermediate  tarsi,  .20  mm.; 
length  of  posterior  femora,  1.25  mm.;  length  of  posterior  tibiae,  almost  1.50 
nmi.;  length  of  posterior  tarsi,  almost  .50  mm. 

Since  writing  the  above  I  have  reared  the  insect  through  from  the 
beginning.    There  are  five  instars. 


510  The  University  Science  Bulletin. 


PLATE  LIII. 

The  genus  Curicta  was  represented  in  the  United  States  by  Curicta  howardi 
Mont.,  described  from  a  single  specimen  taken  at  Victoria,  Tex.,  years  ago. 
Nothing  was  known  concerning  the  biology  of  these  bugs.  The  sketch  of  the 
egg  and  first-instar  nymph  are  therefore  of  interest. 


Wiley:    Biology  of  Curicta. 
PLATE  LIII. 


511 


l^t.    In*ta.T    Cur  (tta.  .  hoiwardr. 


K(^<^     **\     CuT»«i,\aL    . 


i 


THE 

KANSAS  UNIVERSITY 

Science  Bulletin 


Vol.  XIV,  No.  21— October,  1922. 

(Whole  Series,  Vol.  XXIV,  No.  21.) 

ENTOMOLOGY  NUMBER  V. 


CONTENTS: 

Biology  and  Morphology  of  Lepyronia  quadrangularis 

(Say) — Homoptera,  Cercopid.e Kathleen  Doering. 


PUBLISHED  BY  THE  UNIVERSITY 
LAWRENCE,  KAN. 


Entered  at  the  post  office  in  Lawrence  as  second-class  matter. 

9-4522 


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SCIENCE  BULLETIN 

Vol.  XI V.J  October,  1922.  [No.  21. 


Biology  and  Morphology  of  Lepyronia  quadrangularis 
(Say) — Homoptera,  Cercopidse. 

BY  KATHLEEN  DOERING. 

Submitted  to  the  Department  of  Entomology,  University  of  Kansas,  in  partial  fulfillment  of 
the  requirements  for  the  degree  of  Master  of  Arts. 

INTRODUCTION. 

THE  purpose  of  this  paper  is  twofold;  first  to  add  to  the  limited 
data  concerning  the  biology  of  the  Cercopidse;  and  secondly,  to 
contribute  to  the  morphological  knowledge  of  the  family,  and  inci- 
dentally of  the  order.  The  subject  of  this  paper  was  suggested  by 
Dr.  Paul  B.  Lawson.  Since  Lepyronia  quadrangularis  (Say)  is  one 
of  our  most  common  spittle  bugs,  it  was  thought  that  a  study  of  its 
biolog\",  habits  and  morphology  would  be  well  worth  while.  Later 
when  a  review  of  the  literature  revealed  the  meagerness  of  both  bio- 
logical and  morphological  details  concerning  other  species  in  the 
family,  the  necessity  for  this  work  was  magnified. 

By  means  of  the  morphological  studies  the  writer  has  attempted 
to  accomplish  two  things:  First,  since  little  work  has  been  done  on 
the  morphology  of  any  cercopid,  to  give  a  detailed  description  of  the 
external  anatomy,  merely  for  the  morphological  interest  involved; 
and  secondly,  to  contribute  a  small  share,  if  possible,  in  determining 
the  relationships  of  the  families  within  the  order.  A  thorough 
knowledge  of  the  morphology  of  all  families  in  a  given  order  seems 
necessary  to  correctly  determine  their  phylogenetic  relationship.  In 
the  literature,  however,  they  are  usually  determined  on  the  basis  of  a 
comparison  of  certain  special  parts  of  the  body  of  a  single  species 
from  each  family.  But  since  the  species,  or  at  least  the  genera,  may 
vary  within  a  family,  some  having  primitive  head  sclerites  and  spe- 
cialized genitalia,  while  others  are  just  the  opposite,  it  follows  that 
a  fairly  thorough  knowledge  of  the  entire  structure  of  the  body 

(515) 


516  The  University  Science  Bulletin. 

should  be  obtained  before  such  relationships  are  determined.     To 
this  end  this  small  contribution  is  made. 

The  writer  wishes  to  express  her  appreciation  to  all  who  have  as- 
sisted in  this  work:  To  Prof.  S.  J.  Hunter  for  his  interest  and  readi- 
ness to  help;  to  Dr.  Paul  B.  Lawson,  under  whose  direction  the  work 
was  done,  and  who,  at  the  sacrifice  of  his  own  research  time,  gave 
most  helpful  advice  and  criticism;  to  Dr.  Grace  M.  Charles  for  the 
correct  identification  of  most  of  the  host  plants ;  to  Lucy  M.  Hack- 
man,  Dr.  H.  B.  Hungerford  and  P.  A.  Readio  for  their  kindly  help 
and  criticism,  and  to  all  others  who  in  any  way  have  assisted  in  this 
work. 

THE  SYSTEMATIC  POSITION. 

FAMILY  CHARACTERISTICS. 

The  Cercopidse  are  distinguished  from  the  other  families  of  Ho- 
moptera  by  having  three-segmented  tarsus,  two  ocelli,  the  antennse 
inserted  in  front  of  and  between  the  eyes;  the  prothorax  not  pro- 
longed backward,  and  the  hind  tibiae  armed  with  one  or  two  stout 
teeth,  with  two  rows  of  short,  stout  spines  at  the  tip. 

SUBFAMILY  CHARACTERISTICS. 

The  Cercopidse  are  divided  into  two  subfamilies — Cercopinae  Am. 
&  Serville,  and  Aphrophorinse  Am.  &  Serv.  Lepyronia  quadrangu- 
laris  (Say)  comes  within  the  latter.  According  to  Ball  (1898)  the 
Aphrophorinse  are  separated  from  the  Cercopinae  by  the  following 
characteristics:  Anterior  margin  of  the  pronotum  angulate;  head 
equaling  or  almost  equaling  the  pronotum  in  width;  elytra  com- 
pressed behind,  rarely  reticulate. 

HISTORICAL  REVIEW. 

This  insect  has  been  described  under  various  genera.  It  was  first 
made  known  to  science  by  Say  in  1825  as  Cercopis  quadrangularis. 
In  1831  is!  was  put  in  the  genus  Aphrophora  by  Say,  and  in  1851  in 
the  genus  Ptyelus  by  Walker.  Amyot  and  Serville  (1843)  described 
it  under  the  genus  Lepyronia,  which  name  it  now  holds. 

GENERIC  DESCRIPTION. 

The  original  description  of  the  genus  by  Amyot  and  Serville  is 
as  follows:  "Corps  court  et  remasse.  Tete  en  cone  arrendi  anteri- 
eurement,  sans  carene  longitudinale  mediane  sur  le  vertex  ni  sur  le 
front.  Elytres  bombees,  en  ovale  court  et  en  forme  de  coquille.  Les 
autres  caracteres  sont  ceux  des  Aphrophores.  Du  gree  /^-'V'Sr, 
coquille  d'oeuf." 


Doering:    Lepyronia  quadrangularis.  517 

The  genus  Leypronia  is  separated  by  Ball  from  the  other  genera 
of  Aphrophorinse  by  these  characteristics:  Anterior  margin  of  ver- 
tex between  front  and  eyes  sharp ;  ocelli  nearly  equally  distant  from 
eyes  and  each  other;  rostrum  short,  not  exceeding  middle  coxae; 
anterior  margin  of  pronotum  rounded;  corium  without  terminal 
membrane ;  whole  upper  sm-face  densely  pubescent,  hiding  sculptur- 
ing and  venation. 

KEY   TO   SPECIES  * 

A.  Margins  of  the  vertex  regularly  rounding  to  the  obtuse  tip;  elytra 
islightly  angularly  inflated,  nearly  twice  longer  than  their  combined 
width   (folded),  graj-ish,  testaceous,  with  a  distinct  V  on  each  elytron. 

quadrangularis    (Say) . 

AA.    Margins  of  vertex  straight  or  concave,  the  tip  slightly  produced;  elytra 
inflated,    no    more    than    one-half    longer    than    their    combined    width 
(folded). 
B.    Small,    testaceous,    rather    narrow;    the    vertex    broad    and    short, 
shorter  or  only  equaling  the  pronotum  in  length;   apex  of  elytra 
broadly  subhyaline.  angulifera  (Uhl). 

BB.   Large,  nearly  uniform  grayish,  general  form  globose;  vertex  longer 
than  pronotum.  gibbosa  (Ball) . 

The  original  description  of  Say  is  given  herewith: 

Brownish-cinereous  elytra  with  two  oblique  brown  bands  confluent  at  the 
outer  margin;  beneath  black;  feet  annulate  with  pale.  Body  brownish 
cinereous,  covered  with  dense,  minute  hairs;  head  obsoletely  spotted;  eyes 
fuscous,  a  pale  longitudinal  line  on  the  middle,  in  which  is  a  brown  central 
line;  stemmata  indistinct,  black;  thorax  emarginate  at  the  anterior  angles 
for  the  reception  of  the  eyes,  and  deeply  emarginate  behind  for  the  reception 
of  the  scutel;  a  double  series  of  obsolete,  indented  spots  before;  scutel,  tip 
and  basal  angles  acute;  hemelytra  pale  brownish  cinereous;  an  oblique  black- 
brown  fascia  from  inner  basal  angle  is  confluent  at  the  middle  of  the  exterior 
margin,  with  an  oblique  fascia,  which  terminates  near  the  sutural  tip;  tip  with 
a  small  blackish  curve;  region  of  humerus  dusky,  beneath  black;  feet  black; 
thighs  annulate  with  pale;  posterior  pair  of  tibiae  pale,  armed  with  two 
robust  spines  behind  and  numerous  small  ones  at  the  tip;  posterior  tarsi 
armed  with  spinules  at  the  tips  of  the  first  and  second  joints  beneath;  abdomen 
black;  tail  pale  beneath. 

The  following  is  a  technical  description: 

Form.  Length:  9  6.4  to  7.2  mm.;  5  6  to  6.8  mm.  Width:  5  3  to  3.2 
mm.;  $  2.4  to  3  mm. 

Vertex  flat  or  depressed;  length  and  width  about  equal,  as  long  as  pronotum, 
margins  rounding  to  a  bhmt  apex;  tylus  large,  parallel  margined,  nearly  one- 
half  length  of  vertex;  eyes  level  with  vertex,  distended  below;  front  somewhat 
inflated,  rising  gradually  from  the  sides,  margins  convex,  slightly  longer  than 
wide;  clypeus  only  shghtly  inflated,  twice  as  long  as  wide;  pronotum  flat,  twice 
as  wide  as  long,  slightl}'  emarginate  anteriorly  with  transverse  rows  of  im- 

*BaIl,  E.  D.  A  Review  of  the  CereopidsB  of  North  America  North  of  Mexico.  Rept.  la. 
Acad.  Science,  1898. 


518  The  University  Science  Bulletin. 

pressions  behind  margins;  lateral  margins  nearly  parallel,  longer  than  short 
diameter  of  eye;  elytra  not  quite  twice  as  long  as  wide,  outer  margin  flaring, 
apex  angulate;  abdomen  broadly  triangular,  about  as  wide  as  long,  margins 
and  apex  greatly  exceeded  by  wings. 

Color.  Vertex  and  tylus  mottled  testaceous  brown;  minute  yellow  line  on 
margins;  eyes  deep  fuscous;  face  uniform  testaceous  brown;  pronotum  reddish 
brown  at  base,  fading  into  brown  cinerous;  elytra  grayish  or  tawny  brown,  a 
patch  at  the  base,  another  at  apex,  an  oblique  band  from  tip  of  scutellum  to 
a  point  beyond  middle  of  costa,  another  from  point  of  claws,  meeting  this  on 
costa  and  forming  a  V  on  each  elytron,  brownish  fuscous;  abdomen  blackish 
brown,  apex  paler;  legs  testaceous  brown,  annulate  with  pale,  spines  on  pos- 
terior leg  very  black  at  tips. 

LIFE  HISTORY. 

HISTORY. 

Life  history  notes  on  the  Cercopidae  are  rather  limited.  Osborn 
(1916)  made  some  valuable  studies  of  the  life  histories  of  Maine 
froghoppers,  but  gave  no  complete  history,  including  the  egg  stage 
and  five  nymphal  stages,  of  any  one  species.  He  figures  and  de- 
scribes three  instars  of  Lepyronia  quadrangularis.  In  1921  Garman 
published  the  life  history  of  Philcenus  lineatus  (Linn.),  wherein  he 
describes  the  egg  and  four  instars.  Later  Barber  and  Ellis  (1922) 
described  the  oviposition  of  three  species,  Philcenus  lineatus  (Linn.) 
and  Philaroiiia  bilineata  (Say.)  The  most  complete  study  of  a  cer- 
copid  life  history  is  that  of  a  foreign  form,  Tomaspis  varia,  which  is 
a  pest  of  sugar  cane  in  Trinidad  (Urich,  1913).  Garman  (1923) 
gives  a  complete  description  of  the  life  history  of  Clastoptera  obtusa 
(Say)  and  a  brief  description  of  the  egg  stage  of  Lepyronia  quad- 
rangularis (Say). 

DISTRIBUTION. 

Lepyronia  quadrangularis  (Say),  according  to  the  Snow  collec- 
tion, has  been  taken  from  four  counties  in  Kansas,  namely,  Chero- 
kee, Douglas,  Neosho,  and  Doniphan.  It  is  also  recorded  here  from 
Atherton,  Mo.  Ball  (1898)  lists  it  from  Ontario,  New  Hampshire, 
New  York,  Pennsylvania,  Connecticut,  District  of  Columbia,  Mary- 
land, West  Virginia,  Georgia,  Florida,  Mississippi,  Ohio,  Iowa, 
South  Dakota,  Nebraska,  Colorado  and  Texas.  Van  Duzee  lists  it 
from  Muskoka  lake  district  of  Canada,  Lake  Temagami,  Ontario 
and  Quinze  lake  region.  Comstock  says  it  is  one  of  the  most  com- 
mon spittle  bugs  of  eastern  United  States. 


Doering:   Lepyronia  quadrangularis.  519 

HABITAT. 

Lepyronia  quadrangularis  is  usually  associated  with  weed  patches, 
and  therefore  might  truly  be  called  a  weed  insect.  Yet  its  habitat  is 
not  quite  so  general  as  this  would  imply.  Neither  adults  nor 
nymphs  can  be  taken  on  every  weed  patch  one  runs  across  in  col- 
lecting, but  seems  to  be  found  only  in  particular  places.  The  se- 
lected spots  usually  occur  near  woods  or  thickets.  It  has  been  taken 
in  thickets  where  weeds  have  sprung  up  in  open  spaces,  by  the  road- 
sides adjoining  woods  or  thickets,  and  on  the  weeds  growing  at  the 
edge  of  a  wheat  field  and  on  the  wheat  itself.  They  were  found  in 
most  abundance,  however,  in  a  weed  patch  occupying  half  of  a  city 
block.  This  plot  offered  a  variety  of  host  plants  and  shelter,  such 
as  dogwood  and  elm  sprouts,  and  large  patches  of  sweet-clover,  rag- 
weed and  grasses;  in  fact,  most  of  the  collecting  necessary  for  this 
work  was  done  at  this  place,  and  several  thousands  of  spittle  insects 
must  have  been  taken  here. 

HIBERNATION  AND  SPRING  APPEARANCE. 

Ball  (1920)  stated  that  all  Cercopidse  except  one  overwinter  in 
the  egg  stage.  Having  this  idea  in  mind  when  these  studies  were 
started,  it  was  expected  that  the  adults  soon  after  emerging  would 
mate,  the  female  lay  her  eggs,  and  the  overwintering  form  be  easily 
obtained.  The  matter,  however,  was  not  as  simple  as  this.  During 
the  summer  of  1921  adults  were  observed  in  the  field  until  August, 
but  no  mating  or  oviposition  took  place.  The  following  spring, 
toward  the  latter  part  of  April,  eggs  were  sought  in  the  field,  but 
none  could  be  found.  On  April  29  the  first  instars  were  found.  At 
this  time  Mr.  C.  H.  Curran  gave  the  writer  two  adult  females  which 
he  had  collected  on  April  1  and  April  4.  He  stated  that  toward  the 
latter  part  of  March  and  April  adults  had  been  quite  abundant. 
This  evidence  seemed  to  point  to  the  supposition  that  the  adults 
might  overwinter.  Throughout  the  summer  adults  were  closely  ob- 
served in  both  the  field  and  laboratory.  During  August  and  Sep- 
tember adults  were  very  abundant,  an  average  of  fifty  being  taken 
in  an  hour's  collecting.  On  September  27,  seventy-five  adults  were 
taken;  on  October  9,  sixty;  and  October  19,  thirty.  In  one  of  the 
outside  cages  four  adults  were  observed  on  November  23.  These 
late  occurrences  were  due,  no  doubt,  to  the  very  late  season,  which 
prolonged  vegetation  as  well  as  insect  life. 

In  the  late  fall,  after  mating  had  taken  place,  an  experiment  was 
attempted  for  the  purpose  of  obtaining  a  premature  oviposition. 


520  The  University  Science  Bulletin. 

Twenty  adults  were  placed  in  a  glass  jar  containing  sterilized  leaves 
and  twigs  and  covered  by  cheese  cloth.  The  jar  was  then  placed 
outdoors,  surrounded  by  ice  except  for  a  small  breathing  space  at 
the  top  and  kept  in  this  condition  for  eight  days.  After  a  few  days 
the  adults  became  very  sluggish,  scarcely  moving  when  disturbed 
except  for  a  slight  motion  of  the  legs.  When  the  eight  days  had 
elapsed  they  were  brought  into  a  warm  room  and  placed  in  a  cage  in 
which  green  plants  were  growing.  Nine  out  of  twenty  insects  sur- 
vived, but  no  oviposition  occurred.  Although  not  fulfilling  its  orig- 
inal purpose,  the  experiment  seemed  to  show  that  the  adults  probably 
overwinter.  On  December  23  Mr.  Beamer  collected  a  female  of  this 
species  while  sifting  leaves  in  Cherokee  county.  Four  overwintering 
females  were  taken  on  April  18  and  19  of  this  year.  The  exact  place 
of  hibernation  is  difficult  to  determine.  To  date  no  adults  have  been 
obtained  from  the  outside  cage.  They  probably  overwinter  deep 
down  under  the  leaves  and  matted  grass  in  the  cracks  and  crevices 
of  the  ground.* 

CLIMATOLOGICAL  DATA. 

As  was  stated  in  the  foregoing  paragraphs,  overwintering  females 
were  taken  on  April  1  and  April  4  in  the  spring  of  1922,  and  per- 
haps others  could  have  been  obtained  earlier,  while  in  1923  the 
first  ones  were  taken  on  April  19,  making  a  total  difference  of 
seventeen  days.  This  great  variation  can  easily  be  explained  by 
the  difference  in  temperature  of  the  two  years.  A  comparison  of 
the  two  springs  is  given  in  the  following  table: 

Mean  Departure.  Lowest  Highest  Greatest 

19S2.  for  month,    from  normal.      temperature.  temperature,    daily  range. 

February  34.2°  +1.47  2°  on  13th  75°  on  21st  39o 

March    44.5°  +1.4  10°  on  1st  78°  on  23d  36° 

April    56.3°  +1.7  31°  on  1st  82°  on  6th  38° 

ms. 

February    30.3°  — 0.54  —2°  61°  38° 

March    40.74°  —2.03  ^1°  80°  44° 

From  a  study  of  the  table  it  is  readily  seen  that  the  spring  of 
1922  was  above  normal,  while  this  spring  is  below  normal,  which 
accounts  for  the  great  difference  in  the  dates  of  spring  emergence. 

SEASONAL  history  AND  SUMMARY  OF  LIFE  HISTORY. 

A  brief  seasonal  history  and  summary  of  life  history  is  as  fol- 
lows: The  insect  spends  the  greater  part  of  the  year  in  the  adult 
form,  covering  a  period  of  about  ten  months,  and  the  other  two 
months  are  spent  in  the  egg  and  nymphal  stages.  Mating  takes 
place  in  the  fall;  the  females  overwinter,  emerging  in  the  spring 
along  in  April.     A  few  days  after  emergence  the  eggs  are  laid. 


Doering:    Lepyronia  qladraxuularis.  521 

The  egg  stage  probably  lasts  two  weeks,  followed  by  a  six  weeks' 
nymphal  stage.  The  adults  appear  in  June  and  feed  all  summer 
and  fall  until  after  mating,  when  the  males  probably  die  and  the 
females  go  into  winter  quarters. 

OVIPOSITION. 

Females  collected  in  the  spring  were  confined  on  small  Solidago 
and  sweet-clover  plants  under  lamp  chimneys.  Three  females  col- 
lected on  April  18  died  on  April  23.  Two  of  the  females  apparently 
laid  no  eggs.  Thirty-eight  eggs  were  found  in  the  abdomen  of  one 
and  thirty-three  in  the  other.  The  third  female  laid  in  all  nine 
eggs,  one  of  them  being  found  alone  in  one  leaf  and  the  other  eight 
in  a  group  in  another  leaf.  The  latter  were  laid  in  a  row  along  the 
slender  petiole  of  the  Solidago  leaf.  Both  leaves  in  which  the  eggs 
were  inserted  were  brown  and  withered  and  were  found  at  the  base 
of  the  plant.  T